Mammals of the Soviet Union G. Heptner . А. Sludsku The present book, part of a three- volume monograph, 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 information on their external morpho- logy, 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 wildlife painter, А.М. Котагоу, 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. i) ар ИИ, Ой ie Vis ean in Z uy ne Ч ЕЕ АИ ИИ д Зе ibe И ne Te git Mammals of the Soviet Union Volume II Part 2 MAMMALS OF THE SOVIET UNION In Three Volumes Edited by © V.G. Heptner and N.P. Naumov Vysshaya Shkola Publishers Moscow, 1972 Mammals of the Soviet Union Volume II, Part 2 CARNIVORA (Hyaenas and Cats) V.G. Heptner and A.A. Sludskii Illustrators A.N. Komarov and N.N. Kondakov Scientific Editor ROBERT S. HOFFMANN Smithsonian Institution Libraries and The National Science Foundation Washington, D.C. 1992 TT 76-52041 Mlekopitayushchie Sovetskogo Soyuza In Three Volumes V.G. Heptner and N.P. Naumov, editors Vysshaya Shkola Publishers Moscow, 1972 Translator: P.M. Rao General Editor: Dr. V.S. Kothekar © 1992 Amerind Publishing Co. Pvt. Ltd., New Delhi Library of Congress Cataloging-in-Publication Data (Revised for volume 2, pt. 2) Geptner, V.G. (Vladimir Georgievich), 1901-1975. Mammals of the Soviet Union. Translation of: Mlekopitaiushchie Sovetskogo Soitiza. Bibliography: v. 2, pt. 2, p. Vol. 2, pt. 2 has index. Supt. of Docs. по.: SI 1.2: Ar7/2 Contents: v. 1. Artiodactyla and Perissodactyla— Ward: —pt. 2. Carnivora (Hyaenas and cats). 1. Mammals— Soviet Union. Г. Nasimovich, A.A. Il. Bannikov, Andrei Grigor’evich. II. Hoffmann, Robert $. IV. 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 Pauls Press, New Delhi 596.5. G 36 The present book, part of a three-volume monograph, 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 information 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 wildlife painter, A.N. 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. Ра а 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. The present volume of the series Mammals of the Soviet Union, Carnivores (Hyaenas and Cats), is the second volume to be published in English. Previously Volume I, Artiodactyla and Perissodactlya, was published in 1988. A third English translation, Volume II, Part 3, Pinnipeds and Tooth- whales, is in preparation and should be published in the next year or two. This will leave only one part of the original Russian language series untranslated; namely, Volume II, Part 1, Sea cows and Carnivores (Dogs, Bears, and Mustelids). The fact that this latter volume was not published in English translation in the proper sequence is due to the fact that the translation was performed under a different contract from that supervised by P.M. Rao of Amerind Publishing Company, New Delhi, India. However, it is the intention of the Smithsonian Libraries Translation Program to publish this final volume as soon as possible. As indicated in the foreword to the English edition of Volume I, the senior author and editor of the series, Dr. Vladimir Georgievich Heptner, died in 1975, and did not complete the projected series. However, work continues, and volumes on baleen whales and lagomorphs are in progress. It is to be hoped that English translations of these volumes, when they appear, will follow more promptly than has been true of the first volumes of the series. English readers interested in Dr. Heptner’s contributions as a mammalogist 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 Severnaya, Dvina). Soviet administrative units are numerous, and the following equivalents were employed in translation: Krai, territory; oblast’, district; raion, region; Vili 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, I have adopted the following conventions: 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 ап -sk ending this is, however, transliterated (e.g., Omsk); when a Russian ‘‘soft sign’’ is employed in a place name, this is transliterated as ап apostrophe (e.g., Khar’kov). 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. I would appreciate it if readers would bring any errors they may notice to my attention. One further point of confusion not apparent to me when Volume I was translated also requires clarification; that is the English transliteration of the senior author’s surname. This begins with the fourth letter of the Cyrillic alphabet, which usually has a ‘‘G’’ sound in Russian. However, the surname was originally German (as indeed was mine) and in the original German began with the letter ‘‘H’’ of the Latin alphabet. Since Cyrillic has no equivalent of ‘‘H’’ this is usually transliterated into “‘G’’ in Russian. However, I know from conversations with him that Dr. Heptner preferred to use the original Germanic form of his surname rather than the transliterated version, which is rendered as Geptner. The rules of transliteration employed by the Library of Congress do not permit such flexibility, and the attentive reader may notice that Library of Congress cataloguing employs the latter. June 1989 Robert S. Hoffmann Assistant Secretary for Research Smithsonian Institution Washington, D.C. Foreword 5 The present book is a continuation of the second volume of a three-volume monograph on mammals of the Soviet Union and is devoted to a description of terrestrial carnivores of the superfamily Feloidea, i.e., the hyaena (Hyaenidae) and cat (Felidae) families, thus concluding a description of the order Carnivora. In accordance with the scheme laid out in previous volumes, this work also includes a description of the lion, which lived in our country in the historic past. Considerable information has accumulated in the last decade on cats, partly due to current interest in this group. Populations and distribution of these species have rapidly shrunk and some species are on the verge of extinction. This has evoked particular attention in cats not only from a general biological point of view, but also in the context of conserving natural resources. On the whole, information on cats is so extensive that it necessitated their inclusion in a separate book, using a slightly different style of presentation. Part 3 concerns pinnipeds and cetaceans. Since it is preferable to give a description of Soviet marine animals in a single volume, the taxonomic sequence of the description of orders (pinnipeds after carnivores) had to be slightly modified; later, however, the general sequence in the more specialized orders has been maintained (see ‘‘Systematics of the Class Mammalia’’). Within the orders, as in earlier volumes, descriptions of taxa are presented in the order of increasing specialization. All the characters of the groups and species have been given according to the scheme adopted in preceding volumes. A departure has been made only in the case of lions, and the silvestris—libyca, which are quite complex taxonomic examples of semispecies or superspecies. Paleontological data have been briefly summarized, mainly from the works of Simpson (1945) and Thenius and Hofer (1960), and in some cases other more specialized articles. As in earlier volumes. the range descriptions are generalized; only the boundaries are described, based on an analysis of all available data. Only major reference points are indicated for the periphery. As far as possible, historical changes in the range have been traced and, on their basis, ‘‘reconstructed’’ ranges restricted to the historic period established. In spite of all the suppositions involved in such an approach, it is the only way to establish a fairly natural range. A knowledge of the latter is an essential xX prerequisite not only for resolving theoretical aspects of zoogeography, but also for field work for different fauna. The reconstructed range outside the Soviet Union has been treated in very general terms and based on a large number of special studies, but mainly the critically verified accounts of Miller (1912), Sowerby (1923), Anthony (1928), Aharoni (1930), Nezabitovskii (1934), Lukashkin and Zhernakov (1934), Shortridge (1934), Phillips (1935), Heim de Balzac (1936), Allen (1938-1940, 1939, and 1942), Pocock (1934 and 1941), Cabrera and Yepes (1940), Chasen (1940), Hamilton (1943), Harper (1945), Simpson (1945), Anderson (1946), Carter, Hill and Tate (1946), Prater (1947), Tate (1947), Troughton (1948), Anar* (1949), Kalinesku (1951), Roberts (1951), Ellerman and Morrison-Scott (1951 and 1966), Ellerman, Morrison-Scott and Hayman (1953), Bannikov (1954), Laurie and Hill (1954), Miller and Kellog (1955), Kosswig (1955), Siivonen (1956 and 1967), Haltenorth and Trenze (1956), van den Brink (1958), Frechkop (1958), Cabrera (1958 and 1960), Shaw (1955 and 1958), Markov (1959), Misonne (1959), Murie (1959), Hall and Kelson (1959), Hatt (1959), Рис and Tortri¢ (1960), Vasiliu (1961), Burt and Grossenheider (1962), Burton (1962), Cockrum (1962), Atanasov and Peschev (1963), and several others. Special references have not been made to these works in the text. The North American distribution given in the maps, in almost all cases, is based on the data of Hall and Kelson (1959). The morphological characters of the species described are mostly based on the latest data gathered in the last decade in the Zoological Museum of Moscow University. These have made possible a fresh revision of the geographic variation of all species. For parts of the range outside the Soviet Union the list of geographic forms has usually been given without comment, or only with some general remarks. All synonyms, as in previous volumes, have been worked out anew and given in strictly limited numbers on the basis of the following principles: only true synonyms have been mentioned; from among generic synonyms only those for which species of Russian fauna serve as type species, or if the generic name is of importance, or is sometimes used (or was previously), or could be applied to Russian species. All names, prevailing or otherwise, have been given among synonyms of species in the following instances: 1) if the name is from a nominate species from Soviet territory; 2) if the name is based on a nominate species from other territories, but is of importance and is used or was previously for one or another Russian form; 3) if the name is based on animals from other territories and not ‘For more details on the reconstructed range, see V.G. Heptner, Dinamika areala nekotorykh kopytnykh 1 antropokul ‘turnyi faktor (Range dynamics of some ungulates and the anthropocultural factor). Voprosy Geografii, 1960, vol. 48. Moscow. *Nat clear; not in Literature Cited —Sci. Ed. xi applied to Russian forms, but could be of importance for some of them; and 4) the name of the nominate form is invariably given irrespective of its type locality, whether falling within or outside Soviet boundaries. The actual names have not been shown separately among the synonyms of species, but are given in the list of subspecies inhabiting the Soviet Union and the synonyms pertaining to a given subspecies likewise are listed there. Synonyms of subspecies inhabiting regions outside Soviet boundaries have not been given. Synonyms of a group above the genus have not been given and no references are cited for descriptions; these, unless otherwise stated, are from Simpson (1945). The names given to domestic forms, as adopted and clearly explained in Volume I, have not been used for wild species; the name given to the wild form has been used as the species name. The total number of species in the class is about 3,500? and the number of species in Russian fauna about 300 (Heptner, 1956). In fact, recently an intense study of karyotypes in some cases has led to modifica- tions of apparently natural patterns, which were ascertained in the last decade by the morphogeographic method. Studies based on karyosystematics often result in the recognition of an increased number of species. However, at present, only a small number of species of this class have been studied from this viewpoint, mostly isolates of various species. It is therefore still difficult to establish definitely the extent of such an increase. Judging from the fact that whatever has been described is based on special studies of Russian fauna, major revisions relating to the number of species in the class are not likely. In the present volume A.A. Sludskii compiled the sections ‘‘Biology”’ and “‘Practical Significance’’ for all species except the lion. The remaining text has been written by V.G. Heptner, who also compiled all the range maps except those for which compilers have been duly acknowledged. V.G. Heptner is responsible for the general classification, limits, number, and sequence of species; number, limits, and identification of subspecies; and all the synonyms adopted in this volume. Almost the entire literature published up to 1967 and some of 1968 has been reviewed. Only in a few cases are reports from more recent works occasionally cited. Information on the Amur cat, the European forest and steppe cats and the biology of hyaena has been greatly supplemented or elaborated from the latest information of early 1971. Only cited works have been referenced in the Literature Cited. References to literature have generally been placed in parentheses in the text, giving the surname of the author (without initials) and year of publication; initials are given only in those cases when more than one scientist has the same surname. References 2Е. Mayr (1969) estimates 3,700 and Th. Haltenorth (1969) 4,146. ХИ to persons who have provided verbal or written communications of unpublished data are identified, unlike published references, without a date but with the initials of the author. Naturally, both authors of the present volume have extensively incorporated in the text their own unpublished data, especially since cats have long attracted their attention. In addition, various types of informa- tion still unpublished have been kindly furnished by V.K. Abramov (Vladivostok), Yu.V. Averin and M.N. Lozan (Kishinev), N.K. Vereshchagin (Leningrad), G.D. Dul’keit (Stolba preserve, Krasnoyarsk), N.I. Ishadov (Ashkhabad), B.A. Kuznetsov (Moscow), O.N. Nur-Gel’dyev (Ashkhabad), V.E. Prisyazhnyuk (Sudzukhin preserve), Yu.F. Sapozhenkov (Kostroma), V.N. Skalon (Irkutsk), N.D. Sysoev (Vladimir), B.F. Tserevitinov (Moscow), K.K. Chapskii (Leningrad), and many other colleagues, including workers in several regional game farms, who have been duly acknowledged at the appropriate place in the text. С.Е. Bromlei and А.С. Pankrat’ev have provided particularly important data on the Far East. Pankrat’ev also furnished extremely valuable systematic collections. Sketches of animals in both color and black and white, as in the preceding volumes, are by the famous wildlife painter А.М. Komarov. Drawings of skulls and all anatomical sketches are by the zoologist-artist, N.N. Kondakov. They have prepared all the foregoing especially for this book from specimens in the Zoological Museum of Moscow University. Only the drawing of the hyaena skull was made by V.N. Lyakhov. The photographs in the biological sections of this work have been taken in part from other works by V.G. Heptner. Photographs have also been provided by V.K. Abramov (Vladivostok), Yu.K. Gorelov (Badkhyz preserve), S.V. Marakov (Kirov), A.G. Pankrat’ev (Vladivostok), V.E. Prisyazhnyuk (Sudzukhin preserve), and others, whose names have been duly acknowledged at the appropriate place. All photographs, with a few exceptions, are published here for the first time. Moreover, all illustrations are original and have been selected by V.G. Heptner. The constant cooperation of the Chief Librarian of the Zoological Museum of Moscow University, V.I. Korotkova, especially in making available rare publications, is deeply appreciated. ‚ Cordial acknowledgement is given here to all the persons listed above and to others who in any way assisted in the preparation of this work. The authors are particularly grateful to Prof. I.B. Volchanetskii (A.M. Gorkii Kharkov State University) and to Prof. A.P. Korneev (T.G. Shevchenko Kiév State University), who took upon themselves the task of reading the manuscript and offering many useful comments, and to our colleague, I.P. Mitina of the Department of Vertebrate Zoology, who provided extensive and competent editorial assistance. This work was mostly executed within the walls of the Zoological xiii Museum of Moscow University and is based to a great extent on the rich collections held there.* The authors are deeply obliged to the Director of the Museum, O.L. Rossolimo and to all the workers in the Division of Mammalogy. The first two publications under this monograph have been well received in the Soviet Union and abroad. The German translation of Volume I (С. Fischer, Jena, 1966) has also been well reviewed by the international press. The authors hope that this volume likewise will enjoy a good reception. They request all those using this publication to communicate errors and suggestions for improvement to Prof. Vladimir Georgievich Heptner, Zoological Museum of Moscow University, 6 Gertsen Street, Moscow 103 009. *Museum material has been designated by the abbreviation ZMMU (or KZMMU) and that of the Institute of Zoology of the Academy of Sciences of the USSR as ZMAN (or KZMAN). Table of Contents* FOREWORD TO THE ENGLISH EDITION vil FOREWORD ix SYSTEMATICS OF THE CLASS MAMMALIA хх KEY FOR IDENTIFICATION OF ORDERS ххШ ORDER CARNIVORA BOWDICH, 1821 1 Key for Identification of Families in the Order Carnivora 1 Family HYAENIDAE** Gray, 1869 (Hyaena) р Subfamily Hyaeninae** Mivart, 1882 8 Genus Hyaena Brisson, 1762 (Striped Hyaena) 8 Нуаепа hyaena Linnaeus, 1758 10 Diagnosis ill Description 11 Systematic Position 16 Geographic Distribution 16 Geographic Variation 20 Biology 21 Practical Significance 45 Family FELIDAE Gray, 1821 (Cats) 47 Key for Identification of Species of the Cat Family, Felidae 66 Genus Panthera Oken, 1816 76 Panthera leo Linnaeus, 1758 (Lion) 83 Description 83 Systematic Position 86 Geographic Distribution 87 Geographic Variation op) Biology 94 *Pages 550-551 in Russian original. The English table of contents is not a literal translation — Sci. Ed. **Transposed in Russian original —Sci. Ed. Panthera tigris Linnaeus, 1758 (Tiger) 95 Diagnosis 96 Description 96 Systematic Position 106 Geographic Distribution 107 Geographic Variation 129 Biology 145 Practical Significance 193 Panthera pardus Linnaeus, 1758 (Leopard) 203 Diagnosis 204 Description 204 Systematic Position : 2 Geographic Distribution PPD Geographic Variation 228 Biology 233 Practical Significance 267 Genus Uncia Gray, 1854 (Snow Leopard) 274 Uncia uncia Schreber, 1775 276 Diagnosis 276 Description 276 Systematic Position 286 Geographic Distribution 286 Geographic Variation 292 Biology РОВ Practical Significance 314 Genus Felis Linnaeus, 1758 (Small Cats) 319 Felis (Felis) euptilura Elliot, 1871 (Amur, or Far-eastern Forest Cat) 328 Diagnosis 328 Description 329 Systematic Position 335 Geographic Distribution 357 Geographic Variation 342 Biology 346 Practical Significance 355 Felis (Felis) chaus Gildenstaedt, 1776 (Jungle Cat) Diagnosis 7 Description Systematic Position Geographic Distribution Geographic Variation Biology Practical Significance Felis (Felis) silvestris Schreber, 1777 (Wild Cat) Diagnosis A. Group silvestris (European Forest Wildcats) Description Systematic Position Hybridization of Wild with Domestic Cats, and Hybrid Populations Geographic Distribution Geographic Variation Biology Practical Significance B. Group libyca (Steppe Wildcats) Description Origin of Domestic Cats Systematic Position Geographic Distribution Geographic Variation Biology Practical Significance C. Felis (Lynx) caracal Schreber, 1776 (Caracal) Diagnosis Description Systematic Position ХУШ Geographic Distribution Geographic Variation Biology Practical Significance Felis (Lynx) lynx Linnaeus, 1758 (Lynx) Diagnosis Description Systematic Position Geographic Distribution Geographic Variation Biology Practical Significance Felis (Otocolobus) margarita Loche, 1858 (Sand Cat) Diagnosis Description Systematic Position Geographic Distribution Geographic Variation Biology Practical Significance Felis (Otocolobus) manul Pallas, 1776 (Manul) Diagnosis Description Systematic Position Geographic Distribution Geographic Variation Biology Practical Significance Genus Acinonyx Brookes, 1828 Acinonyx jubatus Schreber, 1775 (Cheetah) Diagnosis Description Systematic Position 509 512 513 523 524 526 526 536 538 550 564 631 636 636 636 645 647 650 651 664 665 665 666 674 674 680 681 695 696 702 703 703 709 Geographic Distribution Geographic Variation Biology Practical Significance LITERATURE CITED INDEX OF LATIN NAMES OF ТАХА* *The Russian original also included an Index of Russian Names, omitted here. xix 709 714 Uy VAY 734 VUE Е м а, И _ Systematics of the Class Mammalia 9 The classification of major subdivisions (orders and higher taxa) adopted in the present publication is based on the systematic monograph of the class by Simpson (1945), who extensively relied on paleontological data. It is very close to the system of classification proposed by M. Weber (1928), which in its time was well founded on the anatomy of extant and fossil forms. The orders proposed by these zoologists have long been widely recognized and their grouping into higher taxa as suggested by Simpson, raises no serious objections. Extreme division of the orders is a recent trend. Thus marsupials have been divided into three and even five independent orders (opossums — Didelphida, marsupial martens —Dasyuria, marsupial badgers —Peramelia, caenolestids —Caenolestia, and marsupials, wombats, and kangaroos — Phalangeria), cetaceans into two (toothless whales — Музасосей and toothed whales—Odontoceti), primates into three (lemurs —Prosimiae, simians — Simiae, and tarsiers — Tarsioidea), and so on. All these proposals, however, are not yet well founded. Moreover, among mammalian systematists extreme division of the orders is not generally favored. Neither is it commended here. An order should be primarily an integrating and not a divisive concept. Otherwise the same situation will arise in mammalogy as has already occurred in ornithology, where the concept of an order in some systems of classification has largely lost its significance. As a result the scientific aspect of systematics of vertebrates has been undermined. In the classification of orders under consideration here, only one change has been made, i.e., pinnipeds have been elevated to an independent order; usually they are classed as a suborder of Carnivora. Judging from the systematics of recent mammals, these two groups have diverged and followed different directions of specialization. The isolation of pinnipeds in a separate order is as justified as the isolation of some other orders. Based on already established. views, Lagomorpha are also treated here as an independent order (Weber considers them a suborder of rodents). Hence instead of the 17 orders of Weber and the 18 orders of Simpson, we propose 19. Of these, 10 (56.6%) are represented in our fauna, one of them—-sirenians is now extinct (Steller’s sea cow; see Volume II, Part 1). The main subdivisions within the orders in most cases adhere to Simpson’s scheme. By and large his classification is the most well founded. Nevertheless, significant deviations do occur in the present work, mainly XXii at the level of family and especially at the level of genus. Our concept of genus is much broader than Simpson’s. In our opinion a broader concept of generic divisions conforms better to the structure and principles of natural classification. The classification of cats has consequently undergone a general revision in this book. The systematic classification is given in the chart below. Orders present in our fauna are marked with an asterisk. (V.H.) CLASS MAMMALIA Subclass PROTOTHERIA Subclass THERIA Infraclass METATHERIA Infraclass EUTHERIA Cohort UNGUICULATA Cohort GLIRES Cohort MUTICA Superorder FERAE Superorder PROTUNGULATA Cohort Superorder FERUNGULATA | PAENUNGULATA Superorder MESAXONIA Superorder PARAXONIA Order MONOTREMATA Order MARSUPIALIA *Order INSECTIVORA Order DERMOPTERA *Order CHIROPTERA Order PRIMATES Order EDENTATA Order PHOLIDOTA *Order LAGOMORPHA *Order RODENTIA *Order CETACEA *Order CARNIVORA *Order PINNIPEDIA Order TUBULIDENTATA Order PROBOSCIDEA Order HYRACOIDEA *Order SIRENIA *Order PERISSODACTYLA *Order ARTIODACTYLA ххШ Key for Identification of Огаегз! 1 ( 2). Hind limbs absent; fishlike body with large bilobate caudal fins ЗЕ ВОО а. СЕТАСЕА. 2(1). Hind limbs present; body not fishlike and tail, if present, not in the form of a bilobate fin. 3 ( 4). Forelimbs in the form of leathery wings...... CHIROPTERA. 4 ( 3). Forelimbs of a different structure. 5 ( 6). Fore- and hind limbs very short and resemble paddles, i.e., all the digits entirely enclosed in a common integument cover... = ing SE eG eR St meee a ne en EARP PINNIPEDIA. 6 ( 5). Fore- and hind limbs of a different structure, not paddle-shaped. 7 (10). Feet possess hooves. 8 ( 9). Only one? hoof on each limb....... *PERISSODACTYLA. 9( 8). On each limb, two large hooves and two small ones above them о. * ARTIODACTYLA.? 10 ( 7). Hooves absent (claws present). 11 (14). Between large chisel-shaped incisors and molars on jaws, a large toothless gap (diastema) occurs; its length not less than-length of entire row of molars on corresponding jaw. Canines absent. 12 (13). Two incisors on upper jaw............... RODENTIA. 13 (12). Four incisors on upper jaw; behind each large sharp incisor is located a minute blunt tooth............ LAGOMORPHA. 14 (11). Diastema between incisors and molars absent or much smaller than length of molar row. Canines present. 15 (16). Anterior portion of muzzle extended into well-developed small proboscis. Anteriormost tooth on each jaw, or only upper jaw, much larger than adjacent tooth? .......... INSECTIVORA ‘Orders marked with a single asterisk have been described in Volume I and those with two asterisks partly in Volume II and partly in Volume I.* *No order was double-asterisked in the Russian text— General Editor. ‘This refers only to horses, the only perissodactyls in the Soviet fauna. It may be noted that many perissodactyls have more than one toe on their legs. In fact, their distinctive feature is the odd number of toes—General Editor. Limbs differ in structure in camels. The leg ends in a broadened callused padding and true hooves are absent. There are two highly broadened nails. Camels have not been included in this table as they are domesticated animals (wild camels are extinct). 3If the structure of teeth is different, proboscis is always present and furthermore the front leg extremely short, feet very broad with huge claws, and set on edge with inner surface facing backward (moles). XXIV 16 (15). Anterior portion of muzzle does not form a proboscis. Anteriormost tooth on each jaw not larger than one next to it.......... о а SO Ama KR CARNIVORA.4 4On the coast of the Commander Islands, skulls of the extinct Steller’s cow—a member of the order of sea cows (Sirenia)— have been found. They are distinguished by upper and lower jaws measuring 60 cm in length, in which teeth or traces of them (alveoli) are absent. Steller’s cow has been described in Volume II, Part 1. 15 16 Superorder FERAE Linnaeus, 1758 Order CARNIVORA Bowdich, 1821! Key for Identification of Families in the Order Carnivora” Identification by External Characters 6). Five toes on hind feet. 3). Tail short (shorter than hind foot) and hidden in body fur or barely protrudes. Size very large; body length of adults 1.4 to 2.0 m or О. bears, family Ursidae. 3 ( 2). Tail long (longer than hind foot) and invariably protrudes distinctly from fur. Body length less than 1.2 m. 4 ( 5). Tail distinctly bicolored; six or seven dark-colored transverse rings displayed/omlighter backgrounds... ее. к м raccoon, family Procyonidae. 5 ( 4). Tail monochromatic or terminal part often darker than base, but transverse dark-colored rings absent.................... в en ey em weasels, family Mustelidae. 1). Four toes on hind feet. 8). Four toes on front feet. Neck and back with tall shaggy mane of long resilient hair. Color gray (without rusty tinge) with black stripes traversing trunk....... hyaena, family Hyaenidae (p. 2). 8 ( 7). Five toes on front feet (first toe set above rest). Tall mane along neck and back absent. Coloration different. When trunk covered with black transverse stripes, bright rust-colored tinge of body very vivid. 9 (10). Head elongate, with long muzzle similar to that of a dog or fox. Tail always reaches tarsal joint and is furry, but hair at base short and thinner than in the middle. Claws nonretractile; tips set forward and downward, not highly compressed laterally, and relatively blunt. Coloration without vivid spots or transverse stripes. . . . MAD aC RAT CANTER RAND SOM ay ORIOL SCO wolves, etc., family Canidae. ‘Characteristics of the order have been given in Volume II, Part 1. Families for which the page number is not shown have been described in Volume II, Part 1. 10 ( 9). Head round, with short muzzle like that of a cat. Tail reaches tarsal joint or is shorter; uniformly covered with hair throughout its length, not thinner at base than in the middle. Claws retractile; tips set forward and upward [when retracted], highly compressed lateraily, and very sharp. If claws nonretractile, then body size large (like that of a large dog) and color of body light, with small, sharply defined but not circular black spots. Coat with spots, or transverse stripes, or uniform in color..... cats, family Felidae (p. 47). Identification by Skull Characters 1 ( 6). Palatines extend back beyond transverse line drawn across posterior margin of last teeth by less than one-half their width between those teeth.3 | 2 ( 3). Four teeth [or three] оп each side of lower jaw behind canine; five teeth in upper jaw. Total number of teeth 34 or 32*........ Е. hyaena, family Hyaenidae (р. 2). . Three teeth on each side of lower jaw behind canine; four or three in upper jaw. Total number of teeth 28** ог30............. а. cats, family Felidae (р. 47). 5 ( 4). Seven or six teeth behind canine in lower jaw and six in upper jaw. Total numberof teeth 42 от 90. ois. ee eee а wolves, etc., family Canidae. 6 ( 1). Palatines extend back beyond transverse line drawn across posterior margin of last teeth by more than one-half their width between those teeth. 7 ( 8). Skull large, maximum length exceeds 200 mm............. SEA SOU HONE oe нм bears, family Ursidae. 8 ( 7). Skull small; maximum length less than 200 mm. 9 (10). Six teeth behind canine in upper jaw and six in lower jaw. Total number of ее 40. raccoon, family Procyonidae. 10 ( 9). Less than six teeth behind canine in upper jaw and four, five, or six in lower jaw. Total number of teeth 38 or less....... BS ec ERNE Le Mar ce weasels, family Mustelidae. (V.H.). Superfamily FELOIDEA Simpson, 1931 Family HYAENIDAE Gray, 1869 Hyaenas are specialized, often highly specialized, carnivores of large size. *Error in Russian original; reads ‘‘24’’—Sci. Ed. **Error in Russian original; reads ‘‘38’’—Sci. Ed. ’Measurement should be taken along the suture between the palatines. 17 3 Trunk relatively short, fairly massive, on the whole wolf-like, but rear portion weak and low, withers high, and line of spine slopes down noticeably toward croup. Forelegs high, but hind legs very short. Neck quite short and thick. Head generally resembles a wolf’s, but much larger, massive, and heavier, with a very powerful, relatively short facial portion and heavy lower jaw. Forelegs tall and thin, somewhat bent. Hind legs also appear thin and relatively weak, particularly the thigh. Limbs typically digitigrade; fore- and hind paws with four digits each (digit I absent; in one species five digits occur on forepaws). Digits Ш and IV equal in length, but shorter on average than either digits П or У. Digits with short, powerful, straight, blunt, and nonretractile claws. Digital pads highly developed; digits connected by a dense, thick, and elastic membrane which reaches pad. At base of digits a large callused cushion, roughly semi-crescentoid in shape, occurs. Small carpal pad present high amidst fur on forefoot; pad absent on hind foot. Digital pads bulge notably. Digits always set close to each other (ball-like). Pelage sparse and coarse, with underfur poorly developed or lacking; majority of species have rich mane of long hair running along spine from withers or from head. Color dull gray or brown or rusty-brown, more or less uniform, or with dark (black) transverse stripes or spots. Perineal scent glands absent, but tract (pocket) in anal region with well- developed anal and large sebaceous glands. Skull massive and heavy with powerful facial portion perceptibly extended forward, and compressed, and very small brain case. Viewed from the top, skull appears wedge-shaped with very broad (especially from the rear) and powerful zygomatic arches diverging at an angle. As a result of very powerfully developed sagittal crest, line of dorsal profile from interorbital region to occipital is greatly elevated. Occipital region also highly compressed; when skull viewed from the rear, it has a triangular shape. Sagittal crest forms the extended sharp apex of this triangle. Broadly developed zygomatic arches and compression of brain case associated with development of very powerful masticatory muscles. Alisphenoid canal well developed or absent. External auditory meatus in the form of a tube. Auditory bulla swollen; externally its division into two parts not perceptible; internally, partition wall shifted far back and entotympanic portion very small. Paroccipital processes large and massive, projecting down beyond lower margin of tympanic bulla. Posterior palatine foramina shifted far anterior, to level of second premolar. Dental formula: и и ош 3 1 3 1 1 18 In several respects, dentition is similar to that of the cat family (Felidae). In most species it is very powerful, indeed one of the most powerful in the order, and highly specialized for handling coarse food and crushing bones. Carnassial teeth (upper fourth premolar and first lower molar), especially the upper, very powerful and shifted far posterior to the point of exertion of utmost pressure on the jaws. Inner cusp (protocone) of upper carnassial tooth well developed, shifted to inner surface of anterior end of tooth, and set at a right angle. Other teeth, apart from first upper premolar, and nonfunctional or almost nonfunctional and very poorly developed (rudimentary) upper molar, very powerful with broad bases and cutting edges, but not very sharp cusps. Canines powerful and thick, but not long. As a result of feeding on insects, one genus (Proteles) has highly deviant teeth (see below) and no upper molar whatsoever; this tooth is sometimes absent in the spotted hyaena (Crocuta) also. Vertebral column, including cervical region, of limited mobility. Os penis not developed. Most species are specialized carrion eaters, mostly of ungulates, and sometimes of marine strandings; one species is specialized for feeding on termites and ants. More rarely, they hunt live animals, but mostly prey on the sick and wounded or the young and helpless. They are nocturnal, mostly solitary animals. The den is generally concealed in burrows (mostly those of porcupines), rock crevices, caves, and so on. The geographic range (Fig. 1) is associated with relatively arid regions of the Old World. It occupies all of Africa, from the Cape of Good Hope to the Mediterranean Sea, except for the desert regions of North Africa (Libyan desert and others) and the region of dense humid tropical forests (hyaenas are known, however, in Angola, Togo, Cameroon, and Congo), Arabian peninsula, Asia Minor, Iraq, Iran, Trans-Caucasus, Afghanistan, southern part of Middle Asia, India from the Himalayas to Cape Comorin [Kanyakumari] and in the east as far as Bengal and Nepal-Terai (absent in Burma and Assam). All true hyaenas (striped and spotted—genera Hyaena and Crocuta) are associated in their distribution and attain maximum population levels in regions where ungulates are numerous. The family as a whole represents in all respects a very distinct group, the independence of which is unquestionable. In spite of similarity with wolves in appearance, the affiliation of the family to the superfamily of cats is also clear. The hyaena family occupies a somewhat intermediate position between the relatively generalized family of civets (Viverridae) and the extremely specialized family of cats proper (Felidae). In some respects, hyaenas are closer to the latter. Some species of the family, true hyaenas, are a monotypic and compact 18 ыы 6 Saige Fig. 1. Reconstructed geographic range of the family Hyaenidae. V.G. Heptner. group. However, one genus (Proteles) deviates very sharply. Although a typical representative of the family in its main features, it has several unique features. For example, its dentition is degenerate, associated with its diet of ants and termites: all the teeth are very small, weak, and wide-set; the upper and lower rows together do not form a single cutting apparatus; and most of the teeth do not come into contact with each other when the jaws are closed. The canines and incisors are also as weak as the cheek teeth. Upper molars are absent altogether and the teeth are easily lost during life. Such a dentition cannot serve as an effective ‘‘defensive weapon’’; thus their ability to spray a malodorous liquid from a preanal pouch provides supplementary protection. These characters are important and clearly separate the genus Proteles from true hyaenas (Hyaena, Crocuta). Some systematists regard these features as a justification for placing this genus in a separate family (Protelidae), adjacent to the hyaenas proper—Hyaenidae. The view has also been expressed that Proteles is essentially very closely related, or even belongs to the civet family (Viverridae), constituting a special subfamily within this family. From this viewpoint the similarity of Proteles to the hyaenas, Hyaenidae, is no more than convergence. Such a view has not been accepted, however. It would be proper to regard Proteles as a specialized form of hyaena, evidently separated long ago from the main line of development and hence preserving some characters of viverrids, the family from which hyaenas originated. However, the main thing is that all features of the aardwolf are essentially degenerate features of true hyaenas; there are no major qualitative differences. Hence, systematists of morphological emphasis do not separate the aardwolf into a special family; it is generally allocated to a distinct subfamily. Thus the hyaena family consists of two subfamilies —true hyaenas, Hyaeninae, and the aardwolf, Protelinae. The skull features, especially dentition of the aardwolf as already pointed out, are associated with their specialization for feeding on insects (termites and ants). This feature is also responsible for some other characters, for example, their low mobility. This is one of the very rare, if not unique, instances of extreme specialization of a large carnivore for feeding on insects. In origin, the family is distinctly linked with the viverrids (Viverridae) and may be considered descendant from it. The whole family is relatively young and became distinguishable in the Miocene. The most primitive are the Ictitheriinae, a unique subfamily known from the Lower and Middle Miocene of western Europe to eastern Asia. These are viverrid-like carnivores, usually considered the group ancestral to true hyaenas (Hyaeninae) and the aardwolf (Protelinae). Some, such as Progenetta, are sometimes placed with the Viverridae. However, the Ictitheriinae represent, evidently, only a lateral dying branch of the hyaena family. The aardwolf, Protelinae, among extant animals, is closest to the Ictitheriinae. The remaining fossil forms belong to contemporary groups; all three lineages —spotted hyaena, striped hyaena, and aardwolf—diverged quite early, 1.е., in the Miocene. Spotted hyaenas (Crocuta) were particularly abundant in the Old World. In Europe this group became extinct only in the Pleistocene. Here it was depicted toward the end of its existence as the well-known “‘cave’’ hyaena, С. spelea. The genus Hyaena appeared in the Upper or Middle Pliocene, while the genus Lycyaena (representing the ancestral group) lived in the Lower Pliocene. Throughout its history the family was confined to the Old World and only one species is known from the Lower Pleistocene of North America. 20 й The family is very depauperate. In contemporary faunas, there are only three genera and four species: aardwolf, Proteles Geoffroy, 1824 with one species (P. cristatus Sparrm., 1883); spotted hyaena, Crocuta Kaup, 1828 with one species (С. crocuta Erxl., 1777); and striped hyaena, Hyaena Впз$., 1762 with two species (striped hyaena, H. hyaena Linn., 1758 and brown or shore hyaena, H. brunnea Thunb., 1820). Of the three genera, two (Proteles, Crocuta) wholly belong in Africa and the area south of the Sahara (Ethiopian region), and one genus (Hyaena) with two species (see below) is distributed in Africa (Palearctic and Ethiopian regions), the Near East, and southern Asia. The economic importance of this group is possibly more favorable (scavenging of carrion) than unfavorable, although this differs in some places in accordance with population density, nature of cultivation, etc. Sometimes (southern Africa), hyaenas cause damage to animal husbandry (by killing sheep) and even pose a threat to man. There is only one genus, Hyaena Briss., 1762, found in the Soviet Union. (УН.) 21 Subfamily Hyaeninae Mivart, 1882 Genus of Striped Hyaenas Genus Hyaena Brisson, 1762 1762. Hyaena Brisson. Regn. Anim., 2nd ed., pp. 13 and 168. Canis hyaena Linnaeus, 1758. 1868. Euhyaena Falconer. Paleont. Memoires, vol. 2, p. 464. Canis hyaena Linnaeus, 1758. (V.H.) In general appearance and form the body is wholly typical of the subfamily. Ears large, tall, and pointed. Hair cover relatively long with a large mane of very long, erect, coarse hair along the spine. Color gray with transverse dark stripes on trunk. Newborn with same color pattern as adults. External genitalia of females normal.! Upper molar small but always present and relatively well deveioped; its transverse diameter is approximately three times greater than its length. It is shifted lingual to the tooth row and set at a right angle to the longi- tudinal axis of the carnassial tooth at the level of its posterior margin (as in various species of the cat family, Felidae). The lower carnassial tooth has two large anterior cusps externally [buccal] and a small one internally [lingual], away from the second outer cusp. On the tooth a small but well developed heel occurs, with a tiny tubercle set opposite the upper molar. Unlike the upper jaw, the third and fourth premolars and molar of the lower jaw are almost identical in size, strength, and massive and relatively broad; the carnassial (molar) is equal to the fourth premolar or only somewhat larger than it. All the teeth together form a large and powerful tearing and crushing instrument, occupying much of the dental row, in the region of maximum exertion of force. The outer upper incisors are twice as large as the inner ones. Striped hyaenas are solitary animals, specialized for feeding on carrion (mainly ungulates) and occasionally on marine organisms washed ashore. The reconstructed range of the genus generally coincides with the range of the family. At present striped hyaenas (brown, H. brunnea) are extinct in the southernmost region of Africa. The situation is evidently similar in ‘In the genus Crocuta (spotted hyaena) the structure of the external genitalia of females is altogether different and resembles that of males. The characteristic features of the genus Hyaena have been compared with those of the genus Crocuta. 22 9 some other parts of the mainland, particularly in the northern edge of the range of the brown hyaena in eastern Africa. The taxonomy of the genus (not taking into consideration the geographic variation of the species) is generally fairly well known; the genus consists of just two species: the numerous so-called species of striped hyaena described from Africa and Asia, especially by Matschie and Satunin, have the status of neither species nor race. Of the two genera forming the subfamily of true hyaenas (Hyaeninae), the genus Hyaena should be considered somewhat less specialized than the Fig. 2. Reconstructed species range of striped hyaena, Hyaena hyaena Linn. V.G. Heptner. 23 10 genus Crocuta (spotted hyaena). This is supported particularly by the frequent absence of the upper molar, the general strengthening of the hyaena- type skull, its very large size, the atypical structure of external genitalia (formerly interpreted as attributable to hermaphroditism), the spotted coloration, variation in coloration of newborns and adults, and other features of the spotted hyaena. The genus Hyaena is known from the Middle Pliocene of Europe (Moldavia and western Europe) and from the Pleistocene. Hyaenictis from the Lower Pliocene of Europe and Lycyaena from the Lower and Middle Pliocene of Europe and Asia (China; regarded usually as a predecessor of the former) are closely related to it. The genus Crocuta had already evolved in the Upper Miocene of Asia whence, as also in Europe, it is known until the Pleistocene. Both of these groups of hyaena have survived as independent branches for a very long time (from the Miocene). There are only two species in the genus: the brown or shore hyaena, Н. brunnea Thunberg, 1820, and the striped hyaena, H. hyaena Linnaeus, 1758. Both species are found in Africa. Brown hyaenas are distributed south of the Sahara, as far as the extreme south of the continent (reconstructed range), except for regions of dense forests in western Africa; striped hyaenas occupy the northern region of Africa and penetrate south of the Sahara, mostly in the eastern half of the mainland (see ‘“Description’’ of species). There is only one species in the Near East, Middle Asia, and southern Asia (India). One species, the striped hyaena (Hyaena hyaena Linnaeus, 1758) lives in the Soviet Union and represents about 0.3% of species of the fauna. The range covers the extreme south of the Soviet Union, namely, the Trans-Caucasus and the southernmost regions of the western half of Middle Asia. Economically speaking, partly because of its small numbers, the hyaena has no impact, but is useful in scavenging carrion. Some damage to cattle is not ruled out. Sometimes, apparently, this animal poses a danger even for man. (V.H.) STRIPED HYAENA Hyaena hyaena Linnaeus, 1758 1758. Canis hyaena. Linnaeus. Systema Naturae, 10th ed., vol. 1, p. 40. Benna mountains, Laristan, southern Iran (not India as stated by Ognev, 1931; and others). 1777. Hyaena striata. Zimmermann. Spec. Zool. Geogr., p. 366. Renaming of hyaena Linnaeus. 11 1820. Hyaena fasciata. Thunberg. Sv. Vet. Akad. Handl., vol. 1, p. 59. Renaming of hyaena Linnaeus. 1844. Hyaena vulgaris indica. Blainville. Ostéogr. Mamm., vol. 2, pl. 6. India. 1905. Hyaena vulgaris zarudnyi. Satunin. Izvestiya Kavk. Muzeya, vol. 2, р. 7. Lower course of Karuna River, southeast Iran. 1905. Hyaena bokharensis. Satunin. Ibid., vol. 2, р. 8. ‘Eastern Bukhara’ (southern Tadzhikistan). 1905. Hyaena bilkiewiczi. Satunin. Ibid., vol. 2, р. 9. Ashkhabad region. 1905. Hyaena vulgaris satunini. Matschie. Sitzungsber. Ges. Naturf. Freunde Berlin, p. 363. Trans-Caucasus. (V.H.) Diagnosis H. hyaena L. is the only species of the genus inhabiting the Soviet Union. Description At first sight, the hyaena seems to resemble a wolf or large dog, but possesses an extremely typical external appearance (Fig. 3). Trunk fairly massive but short and set on long legs. Forelegs significantly longer than hind ones, as a result of which the body is raised at the withers, but the croup significantly lower, and the back slopes down. The high withers are the result not only of the difference in leg length, but also intensity of growth of the spinous processes of the corresponding vertebrae, and of the mane running along the back. Legs relatively thin and weak; forelegs perceptibly bent in carpal region. Head placed on thick and fairly long but poorly mobile neck; heavy and massive, with a small, noticeably blunt facial portion. Eyes rather small. Ears very large, broad, set high, and pointed at the tip; wide-set, directed sharply upward (do not ‘‘Агоор”” sideways) and incline somewhat forward. Paws have bulky pads, characteristic of all hyaenas, and powerful but short, blunt claws, which appear trimmed off at the tips. Tail not long and terminal hairs do not descend below calcaneal joint. For an animal of this size, the winter coat is quite long and fairly uniform, with narrow but luxuriant mane of long resilient hair along back from occiput to base of tail. Mane usually ‘‘falling’’ but, on occasion, hairs are capable of standing more or less vertically when erected. Tail also covered with long hair, which appears shaggy. Coat generally coarse, in places bristly, and varies greatly in different seasons. In winter, coat fairly dense, relatively soft, with well-developed underfur and long hair. Guard hairs on trunk (sides) are at this time 50 to 75 mm long, mane hairs 150 to 225 mm, and tail hair about 150 mm. The summer coat is 12 W Gite т bi) Ki Hi ih “le Nps 1) | [РТ КО ‘ tt nr Way Nhe i Ke a fy te RS ON Fig. 3. Striped hyaena, Hyaena hyaena Linn. Sketch by A.N. Komarov. very coarse, much shorter than in winter, sparse, and underfur is absent; the mane is nonetheless large. General shade of winter coat usually dirty brownish-gray or dirty gray. Hair of the mane wide, with pure black or light brown hair terminally and nearly as wide with light gray or almost white hair at base. Small dark bands alternate with much lighter colored sections in basal half of mane; as many as four such bands may occur (Fig. 4). Muzzle, up to eyes and chin, dark, grayish-brown, brownish-gray or black, while top of head and cheeks are lighter in color, usually dirty or grayish-brown. Ears very dark-colored, almost black. Inside as well as outside of ears covered with sparse fur, the color of which depends on skin color. Along edges of ears, hair somewhat more dense and white in color. On throat and downward from front of neck, large black spot separated from chin by a light zone. From flanks, dark field ascends to rear of cheek; some separate 13 black spots occur below dark field. Similar dark spots scattered directly in front of shoulder; individual spots and short dark band located in mane on withers, and fairly long band extends forward and downward from mane through shoulders. Inner and outer surface of forelegs, almost as far as carpus, covered with small dark spots and transverse stripes. No dark patch on tail tuft; tuft usually lighter in color than tail itself. Four indistinct dark vertical stripes, or rows of diffused spots, or a combination of both, seen on flanks. Outer surface of thigh with three or four distinct vertical or oblique dark bands, merging into transverse stripes in lower portion of legs (to heel). Only spots present on legs below heel. Dark stripes and spots brownish-black. Belly dirty gray with a small number of pale spots; latter also present on inner side of thighs. Hair on upper part of tail with same color pattern as mane hair, i.e., black tips but no transverse dark-colored zones in light-colored part at base. Tip of tail black but underfur white. The coloration described above is subject to considerable individual variation with respect to general background, intensity of dark markings, often number and disposition of spots, distinct separation of bands, and other features. The general background varies from fairly dark to rather brown, or straw-colored, or even almost dirty white. Pelage color changes Fig. 4. Striped hyaena in winter coat. Badkhyz preserve, southern Turkmenia. December, 1961. Photograph by Yu.K. Gorelov. 26 27 14 greatly depending on wear and fading; obviously, the color is brighter when the coat is new. Before and during molt the coat is especially dull and pale; at this time the dark shades in particular often fade to brown and cinnamon. Sexual dimorphism in color is absent but seasonal color dimorphism is noticeable. In the short summer coat the stripes and spots are a deeper black, the bands more distinct, and the color contrast far sharper. Newborn young have a silky coat devoid of mane or long hair on the tail. The general basic color is white, muzzle gray, with a broad, dark band extending along the spine, and disappearing on the withers and sacrum. The tail is wholly white. From the shoulders to the sacrum, however, eight vertical black stripes are quite distinct; stray spots between these stripes are likewise well defined. A black stripe extends along the top of the neck (Indian animals; Pocock, 1941). A fairly large pouch without hair is located at the anal opening of males, and large anal glands open into it at some distance above the anus. Several large sebaceous glands also appear between the openings of the anal glands and above them. Scrotum small, covered with hairless skin, and situated in the bare region surrounding the anus. Penis long and preputial folds set far forward. External genitalia of females normal; vulva located in hairless - preanal region, from which it is separated by a narrow strip of hair-covered skin. Large anal glands open into preanal area above anus (Pocock, 1941). There are three pairs of teats. The major skull features are wholly typical of the family and genus, in particular, development of very high sagittal crest; height of crest at the auditory bullae constitutes about one-half the condylobasal length of skull, as does height of nasal region (together with lower jaw). Muzzle short and broad, facial region almost equal to that of brain case or shorter. Postorbital processes well developed, and have rough surface. Frontal region between them inflated (Fig. 5). Anterior and lower margins of orbits bounded by ridge. Zygomatic arch with well-developed postorbital process, and together with supraorbital process of frontal forms posterior, upper, and lower borders of orbit. Orbit closed to a significantly greater extent than in other Russian carnivores (except cats), though not completely. Tip of anterior processes of frontal usually extends to posterior process of premaxilla. Anterior end of nasals deeply notched. Tympanic bullae high (bulging), sharply projecting downward, and constricted anteriorly; their axes lie at an angle to each other. Palate long and broad; distance between posterior ends of carnassial teeth more than one-half length of palate. Deep depression (place of attachment of powerful muscles) occurs on angular processes of lower jaw. Paroccipital processes very large and massive. ys ` OD NS tat . \ АХ ны 7 Gs ey a AAA. © S (д а “hae JSS а к Ч ы 4 Wtf PE С М 42% TO = HEH, SON Rye [С ( NEY Oe SEES NY VRS “is SS: ИЕ СУ, 5 > „ st > У eC 4 \ We M а } ИМ А (У т 7 = \ Е = Z сх MM. <= G i> a с OS 6 Jae 26 Fig. 5. Skull of striped hyaena, Hyaena hyaena Linn. Sketch by V.N. Lyakhov. 16 Features of dentition have already been discussed under generic description. wes Measurements “‘in the flesh’’ of Russian hyaenas are not available. Indian hyaenas (five adult males and five adult females; from Pocock, 1941) are as follows: body length, males, 99.0-109.5 cm, females, 95.5—108.0 cm; tail length, males, 27-34 cm, females, 27—30 cm; length of hind foot, males 20.0—21.5 cm, females, 19.0-22.5 cm; length of ear, males, 15.0-16.5 cm, females, 13.7-15.8 cm; height at shoulders, male (1), 75 cm, female (1), 65 cm. Condylobasal length of skull in males (9), 210-220 mm*, females (9), 202—220 mm*; zygomatic width, males, 149-164 mm, females, 147-155 mm; interorbital width, males, 47—50 mm, females, 44—55 mm; postorbital width, males, 35-39 mm, females 35—40 mm (India; Pocock, 1941). Among hyaenas of Middle Asia (9) and Trans-Caucasus (1) without sex differentiation: condylobasal length, 204—224 mm; zygomatic width 143.0-160.2 mm; interorbital width, 44.1-51.7 mm; postorbital width, 32.2-39.9 mm; mastoid width, 74.1-90.2 mm (Ognev, 1931; material from Zoological Museum, Moscow University). Condylobasal length of an adult female from southern Turkmenia (Pul’-i-Khatum on the Tedzhen) was 221 mm, zygomatic width 157.7 mm, interorbital width 45.9 mm, postorbital width 36.5 mm, and mastoid width 81.8 mm (V.G. Heptner). Males weigh about 38.5 kg, females about 34.0 kg (India; Pocock, 1941). There are apparently no significant sex-related size differences. (V.H.) Systematic Position The striped hyaena is closely related to other species of the genus, the African brown or shore hyaena, H. brunnea. The skull of the latter is larger and its dentition more powerful than in this species; because of these differences the brown hyaena may be considered a somewhat more specialized form. Its body hair coat is very long, the mane less distinct, color dark, and stripes less prominent. (V.H.) Geographic Distribution In open, mainly arid expanses of India, the Near East, and Africa except the southern half. Geographic Range in the Soviet Union The range (reconstructed) in the Soviet Union covers the extreme south *Error in Russian original; reads *‘cm’’—Sci. Ed. 17 of the country, in the Caucasus and Middle Asia, and consists of two separate sections — Caucasian and Middle Asian. These are continuous to the south outside the USSR. The northernmost edge of the range of the species as a whole is located in the Soviet Union (Fig. 6). In the Caucasus the range occupies the plains, foothills, and low mountains of the eastern Trans-Caucasus and displays a complex outline. It includes all of the montane part of Araks valley and the adjoining low hills (hyaenas have been sighted at Merg and Nakhichevan), in the Yerevan depression west of Echmiadzin, and as far as Surmala on the Araks, southwest of Echmiadzin. Hyaenas are distributed farther in the Mugansk steppe, Sal’yansk and Lenkoransk lowlands, and in the Talysh mountains, where the animals have been encountered at altitudes 2,000 m above sea level. In the north the range of the hyaena covers the area along the Alazan, Iori, and Kur’, including Shirak (between Iori and Alazan) and Adzhinour (left bank of the Kur’ below the mouth of the Alazan) steppes (Sheklinsk and Kartalinsk plateaus) and extends to the Karayaz forest (slightly east of Tbilisi) and almost to Tbilisi, and occupies the semideserts in the region of the right tributaries of the Kur’ to the south of Akstafa (Idzhevansk and Shamshadinsk regions of Armenia). The northern boundary also encloses the foothills of the Great [Caucasus] range and extends from there southwest to the foothills and lower belt of the Karabakhsk range (Agdam and other places). Fig. 6. Reconstructed range of striped hyaena in the Soviet Union. V.G. Heptner. 29 18 The Trans-Caucasian section of the range occupies the Apsheron Peninsula and extends in the form of a narrow projection along the coastal region of the Caspian Sea northward to the district between Derbent and Makhachkala— in any case to the lower reaches of the Samur or slightly farther north. Hyaenas extended as far as Akhta along the Samur. Mainly stray animals have been reported along the Caspian coast, possibly due to peculiar circumstances (military activities in the first half of the last century). A reported case of wandering as far as Ordzhonikidze (Vladikavkas) appears dubious. Reports of intrusions to Novorossiisk are undoubtedly erroneous. In Middle Asia hyaenas have been sighted along the Atrek, evidently not reaching the sea, however; in the desert north of the river (Messeriansk desert), and throughout the Kopet-Dag, its foothills and temporarily to the adjacent northern plains, but never penetrating the highest parts of the mountains. In the Great Balkhan range, hyaenas are presently absent but may have lived there in the past. Definite information about this is absent, however. Information about occurrences along the Uzboi (Dement’ev, 1955) is extremely dubious. To the east the range occupied the Tedzhen valley northward, evidently to the city of Tedzhen (possibly even lower), the Murgab valley, evidently to its lower reaches, all of the expanse between the upper divides separating the Murgab and Tedzhen (Badkhyz and Chengurets mountains), the Karabil district lying south between Murgab and Amu-Darya, and southeast Karakum at least to the Karakum canal in the north (villages of Zenkush, Neder-Belent, Kert-Kuyu, Aitysh-Kuyu, and others). Thus, in southeast Turkmenia hyaenas reached deep into the desert, albeit usually along their edges. Along the right bank of the Pyandzh and upper Amu-Darya, the range occupies a fairly significant section of the plains, foothills, and low mountains from the Amu-Darya in the south to the foothills of the Gissar range in the north. In the east the range is bounded by the foothills of the Darvaz range and its southern spurs. The boundary evidently runs slightly east of 70° or along it through the Kulyab, Muminabad, Bal’dzhuan, and Sarykhosor regions. From there, along the foothills of the Gissar range, it turned west, initially through the eastern part of the Gissar valley (region of Ordzhonikidzeabad), then through the Dushanbe region and B. Oktyabrask to Syry-Assiya in the upper course of the Surkhan-Darya, and probably even intersected the valley of this river slightly more to the north. From there the boundary encompassed the southeastern foothill spurs of the Gissar range and Kugitangtau and extended toward the Amu-Darya to the south of this mountain massif—at Kelif. Precise data on temporary incursions of hyaenas along the Amu-Darya pertain to the Termez region and the lower reaches of the Shirabad-Darya. In the Amu-Darya valley, hyaenas have been reported 30 19 from Chardzhou and Farab. These intrusions probably occurred from the south along the river valley. The natural range described above for Middle Asia remains more or less the same today. In the Caucasus, however, mainly before the twentieth century and more particularly in the last 30 to 40 years, it has fluctuated considerably; in fact, hyaenas almost disappeared completely from this area. Even in the 1930s they were encountered in small numbers only in some poorly inhabited pockets of western Azerbaidzhan, eastern Georgia (then and later), the Kartalinsk plateau (in the region of the city of Udabno, southeast of Tbilisi), and probably Talysh. In the 1950s (F.F. Aliev) some stray animals (total of four) were caught in Azerbaidzhan at Yevlakhsk (on the Kur’), Agdzhabedinsk (Mil’sk steppe), and Gadrudsk (southern Karabakh range) region. In 1968 some hyaenas were encountered in the desert hills of Adzhinaur steppe (east of the lower Alazan), Kartalin and Iori plateaus, the Araks valley within Nakhichevan Autonomous Soviet Socialist Republic, and single individuals around Shemakha and in Karabakh (B. Agdamsk and Fuzulinsk regions). According to other data (Alekperov, 1966), hyaenas were absent in southwest Azerbaidzhan even in the early 1950s. Evidently this animal was very rare in the Shirak steppe (1969, Tsiteli-Tskara; A.B. Arabuli). The total disappearance of the hyaena from the Trans-Caucasus is apparently a recent phenomenon. Intrusions from Iran into the extreme south of the Trans-Caucasus can still be expected, however. Extensive hunting only partially explains the disappearance of the hyaena (in the Caucasus hyaenas are accused of carrying off children). More likely reasons would seem to be overall changes in weather conditions, reduction and often disappearance of ungulates, mainly goitered gazelle and in part large predators, and changes in the character of animal husbandry.” According to some reports (Pidoplichko, 1951) contemporary species, or forms close to them, were widely distributed in the Quaternary Period in Europe. Remains have been reported from Ireland, England, Belgium, France, Spain, Portugal, Italy, Switzerland, Germany, Hungary, Czechoslovakia, and Poland, and in the USSR, in the Crimea, Ternopol, Dnepropetrov, and Odessa districts, around Kiev, in Tatariya, the Kuban, and in the southern Urals. In Asia, outside the present-day range, Pleistocene hyaenas are known from the regions of Minusinsk, Krasnoyarsk, the Altai and Trans-Baikal regions, the Mongolian People’s Republic, and China. However, most of these references evidently pertain to cave hyaena *Range based on works of Bogdanov, 1873; Dinnik, 1914; Satunin, 1915; Sultanov, 1930; Ognev, 1931; Leviev, 1939; Heptner and Formozov, 1941; Vereshchagin, 1942, 1947, 1959; Dal’, 1954; Sekunova et al., 1956; Нершег, 1956; Chernyshev, 1959; Nur-Gel’dyev, 1960; Ishunin, 1961; Alekperov, 1966; and others; and unpublished data of V.G. Heptner and F.F. Aliev. 20 (Crocuta spelaea), and fossils of the striped hyaena are known only from Paleolithic caves in Palestine, Morocco, Algeria, and Portugal. Pleistocene finds in France are less reliable; the direct association of the Pleistocene hyaena of England with present-day species is also dubious. According to these data (Vereshchagin, 1959), remains of striped hyaena have not been found anywhere in the Soviet plains, the Crimea, the Caucasus, and Siberia; fossils from the Paleolithic period have only been discovered south of Samarkand. Geographic Range outside the Soviet Union The range (reconstructed) in Asia occupies the Arabian peninsula, including its pre-Mediterranean part, Asia Minor, Iraq, Iran, Afghanistan (except the highest parts of the Hindu Kush), Baluchistan, and all of the northern Indian subcontinent to lower Kashmir, Nepal, and lower Bengal. In the east the range does not include Assam, Bhutan, and Burma. Southward it reaches the Nilgiri hills or even Cape Comorin [Kanyakumari]. Hyaenas do not occur in Ceylon [Sri Lanka]. In Africa the striped hyaena was found, except deep in the central parts of the Sahara and Libyan deserts, in the northern part of the mainland, and south of the Sahara commencing from Azbine (around 19° М. Lat.), in the Sudan, Ethiopia (Abyssinia), Somali, Uganda, Kenya, and northern Tanganyika. At present, the hyaena is already extirpated in some of these areas (Egypt) (Fig. 2). (V.H.) Geographic Variation Geographic variation of the hyaena is insignificant in spite of the fact that a fairly large number of forms have been described (about 24 names). In any case the species and subspecies described outside the Soviet Union, India, and Iran are not justified; only one form exists throughout the whole of India and much, if not all, of the Near East. Its differences from other forms, mainly congruent races, have not been satisfactorily explained and it is tentatively considered the nominal form. In the Soviet Union, therefore, only one subspecies is recognized. Asiatic striped hyaena, H. h. hyaena Linnaeus, 1758 (syn. striata, indica, zarudnyi, bokharensis, bilkiewieczi, and satunini). The description given above pertains to this form. This subspecies is found in the Trans-Caucasus (now almost extinct), southwest and southern Turkmenia, and plains and slightly elevated regions on the right bank of the Pyandzh and upper Amu-Darya from the Darvaz mountains to Kugitangtau. Outside the Soviet Union this subspecies lives in the Asian part of the 31 21 range, except the southern part of the Arabian peninsula (form syriaca apparently does not differ from the nominal form). ok + # The following forms are usually recognized outside the Soviet Union: 1) Н. h. syriaca Matschie, 1900—Syria; 2) Н. h. sultana Pocock, 1934—-southeastern part of the Arabian peninsula; 3) Н. h. vulgaris Desmarest, 1820—Egypt and North Africa to eastern Algeria; 4) Н. h. barbara Blainville, 1844—-western Algeria and Morocco; and 5) Н. h. dubbah Meyer, 1791—Sudan, Somali, Kenya, and northern Tanganyika. The geographic variation of the species evidently requires revision. (V.H.) Biology Population. In the western part of the range within the Soviet Union at the end of the last century, hyaenas were extremely rare on the southwest coast of the Caspian Sea and nearby mountains, extending north to Derbent and Deshlagar (Poletika, 1911; Dinnik, 1914; Heptner and Formozov, 1941). In the eastern and southern Trans-Caucasus, this carnivore was often encountered, but very sporadically. In the 1860s it was ‘“‘common’’ along the Kur’ valley and the adjoining low mountains westward to Tiflis (Tbilisi). In those years hyaenas were caught quite frequently around Tiflis, Yerevan, and Nakhichevan (Radde, 1899). In the Makhats mountains near Tbilisi hyaenas were considered to be “‘extremely common’’ in the 1880s. They were also common at that time in Karayaz. Early in this century the population in this region decreased drastically, and hyaenas disappeared, for example from the vicinity of Tbilisi, but were still ‘‘quite common’’ in the Shirak steppe and Geok-Chai, Echmiadzin, and Surmalin districts. Stray animals were encountered in Zangezur, around Ordubat, near Sal’yan, in the lower course of the Kur’, and in the Talyshinsk mountains along the boundary with Iran (Dinnik, 1914; Satunin, 1915). Commencing in the 1930s the hyaena population in the Trans-Caucasus continued to fall rapidly and the animal disappeared in many regions. Individual animals survived only in the Shchekin and Kartalin plateaus in the Shirak and Adzhinaur steppes (Vereshchagin, 1947, 1959), and in several other regions in the eastern parts of Georgia and Azerbaidzhan. In 1959 hyaenas were caught in Azerbaidzhan in the Burun-Guvakh mountains in Safaraliev region, where they had never been seen before (Episkoposyan and Babakevkhyan, 1959). In the Armenian Soviet Socialist Republic in the first half of the twentieth century, the hyaena was encountered very 22 rarely. In 1925 it was observed in the neighborhood of Navruzl (Artashat region). Later, a single animal was caught in Shamshadinsk region (Dal’, 1954). By 1970 in the Trans-Caucasus the striped hyaena was still encountered singly in the Adzhinaur steppes, in the Kartalin and Топ plateaus and also in the Araks valley in the Nakhichevan Autonomous Soviet Socialist Republic. Sightings and captures were considered a great rarity at this time in Gobustan around Shemakha and in Karabakh (Agdamsk, Agdzhabedinsk, Zhdanovsk, Fazulinsk, and Dzhabrail’sk regions) (Aliev, 1971). Instances are known of their appearance at other places. In 1962 a hyaena was met with in the foothills near Geok-Chai; in 1964 in Nakhichevan Autonomous Soviet Socialist Republic between Khanagya and Kazanchi villages; in 1967 in Adzhinaur steppes, in the Nukhinsk region, and the environs of Kichik- Dakhna village; and in the summer of 1969 in Talysh, Zuvanda, and near the village of Gosmal’yan, Leriksk region. Solitary animals are still sighted in these places from time to time (Kh.M. Alekperov). Even in the past the extreme rarity of the hyaena was noted in Talysh and Zuvanda (Burchak- Abramovich, Mamedov, 1966). The total population of the hyaena in the Trans-Caucasus in 1968 was placed at less than 150 to 200 animals (Aliev, 1971). In eastern Georgia they are ‘оп the verge of extinction’’ (Arabuli, 1970). In the Trans-Caucasus 26 hyaenas were caught from 1930 to 1940, and not more than 5 or 6 from 1940 to 1950 (Vereshchagin, 1959). From 1951 through 1958 on the territory of Azerbaidzhan only four hyaenas were caught, of which two were from Yevlakhsk region, one from Agdzhabedinsk, and one from Gedrudsk (Aliev, 1971). In the last 12 years no skins of this predator have been received in the processing units of Azerbaidzhan, although some animals were caught during those years (Kh.M. Alekperov). In southwest Turkmenia small numbers of hyaenas used to be sighted on occasion in the Atrek valley, and less often along the Caspian Sea coast. For example, near Kara Degish and Chalayuk the hyaena has been reported only a few times in the region of Lake Bol’shoi Delil and Kara-Bab and Kayalydzh mounds. Sightings occurred regularly in these places until 1913, were again reported between 1917 and 1922, and then not until 1941. Near Gasan-Kuli two hyaenas were caught in 1938 and one in 1941. One or two animals were caught at the same place in subsequent years also. The hyaena was seen slightly more often on the Messeriansk plateau (north of the lower Atrek; Dement’ev, 1948, 1955; Samorodov, 1953). In the Kopet-Dag and its foothills the hyaena was not rare but was encountered everywhere, going out also on the adjacent plains. Reports of some nineteenth century researchers (Radde and Wal’ter, 1889; Zarudnyi, 1890; Varentsov, 1894) about the rarity of hyaenas in this region hardly 32 23 reflect the actual situation or indicate а temporary phenomenon. In the first two decades of this century the animal was commonly encountered around Ashkhabad itself, in Gindovar gorge, and in Bagir village (information given to V.G. Heptner by S.I. Bil’kevich). In the 1920s, 1930s, and even 1940s it was common enough, though numbers were not great; in all of the Kopet- Dag mountains and, perhaps, from this area, the predator found its way into Uzboi. At the end of the 1920s a hyaena lair was excavated in the foothills east of Ashkhabad, almost on the plains (V.G. Heptner). The hyaena was rare in much of the territory of the Murgab basin, especially in the northern part (Flerov, 1932). At the end of the 1930s and in the 1940s it was, however, quite common on the upper Murgab and along the Kushka, and some skins (about 10 in 1942) were received in Sary- Yazy for curing every year (V.G. Heptner). In the 1950s the animal became relatively common even in the north. For example, a hunter from the Bairam-Aliisk region caught 18 hyaenas in a few years and 4 in the winter of 1957-1958 (Kleshchinov, 1958). In the extreme southeast of Turkmenia, at Badkhyz, the hyaena was ‘а quite common animal”’ until recently. It was often sighted even in regions adjoining Badkhyz on the north, northeast (Karabil’ upland), and east. Several were caught every year in the environs of Kala-i-mor on the Kushka River. The hyaena was likewise common in the mountains of Gyaz’-Gyadyk at Tedzhen and along the Tedzhen at Serakhs as well as above and below it (Heptner, 1956). The hyaena had become quite rare by 1962 in the low hills of Badkhyz and adjoining regions: during daily excursions for 47 days the predator was sighted only once. Six of the dens discovered were uninhabited (A.A. Sludskii). In the low mountains of Gyaz’-Gyadyk, however, this animal was not particularly rare (Yu.K. Gorelov). By 1970 the hyaena had become rare even in the eastern part of the Karakum close to Karakum canal, where it had persisted at several places in the 1940s and 1950s [Zenkush (Lengych), Neder-Belent, Kert-Kuyu, Aitysh-Kuyu, and others], in fact, it was ‘а commonly occurring”’ large predator (Sekunova et al., 1956; Nur-Gel’dyev, 1960). Over the last 12 years (to 1971) in Turkmenia, only one hyaena skin has been cured although many animals were caught; skins were not sent for curing because they fetched a very low price. On the whole, in Turkmenia during the last 40 to 50 years, the hyaena population and its fluctuations depended on the population of ungulates, primarily arkhar sheep and goitered gazelle, and wolves, who killed these ungulates. The disappearance of goitered gazelle and reduction in the population of mountain sheep in the last decade have determined the fate of the hyaena (V.G. Heptner). The hyaena was rare even in the 1930s to the 1950s in the valleys, a3 24 foothills, and low mountains of the ranges of southwest Tadzhikistan (Flerov, 1935; Chernyshev, 1958). In the 1960s the population in this region decreased even more. During nocturnal observations on predatory animals at water holes and their habitats for over three years, the hyaena was encountered only twice in the Karshitau (Koktau) range at Koktash in the Leninsk region (Arutyunov, 1964). A hyaena was sighted in September, 1969, on the northern slope of this same range (Sambuli), and on its southern slope (Lyaur) in September, 1970 (A.K. Kovalev). In 1967, a young female was caught in the Babatag range (Isambai) (АЛ. Kirkhadzhi). Hyaenas were sighted a few times in the Garua-Niushti range in‘September, 1965, 1966, and 1967 (Dagana-Kiik, Isambai; A.K. Kovalev). In Tadzhikistan not more than 5 to 12 hyaena skins were cured annually from 1932 to 1949. Between 1949 and 1960 three hyaenas were caught alive (D.P. Dement’ev). From 1953 through 1963 only five skins were cured; the animals had been caught in Ordzhonikidzeabad (1955), Kuibyshev and Dangarinsk (1960), and the Pyandzhinsk regions (1963). No skins were received for curing from 1964 through 1970. In Uzbekistan the hyaena was rare in the lower course of the Shirabad River, in Shirabadsk region, although it probably occurred in the adjoining Termezsk region in ‘Чагое numbers on Amu-Darya shore’ (Sultanov, 1939); the latter is extremely dubious. The hyaena was rare along the Surkhan- Darya and in the adjoining mountains. Farther north and eastward, it was absent. By the 1950s the hyaena had become ‘‘а very rare, almost extinct’’ animal in Uzbekistan (Ishunin and Lustin, 1959). Evidently the hyaena is fewer in numbers today in most places in southern Turkmenia and in regions adjoining Afghanistan and Iran. In the Trans-Caucasus, southwest Tadzhikistan, and southeast Uzbekistan, the hyaena is rare and its population has rapidly declined over the years. Soon it will be extinct, especially in the Trans-Caucasus and Uzbekistan. Outside the Soviet Union the hyaena is quite rare in the northern parts of Iran and Afghanistan. It is still common in Baluchistan and numerous at some places in Hindustan (Pocock, 1941; Prater, 1965; Lay, 1967). In Africa the striped hyaena is now rare in northern areas, i.e., Egypt, Libya, Tunisia, and Morocco. It is absent altogether in the Sahara but more common in the savannas of central Africa; it is quite common at some places in eastern parts of Africa (Sudan, Somali, Kenya, and others). In central and east Africa the striped hyaena inhabits the same regions as the spotted hyaena (Crocuta crocuta), but at places where the latter is common, the former is rare, and vice versa (Ronnefold, 1969). Habitat. In the Trans-Caucasus the hyaena inhabits sections of hilly plateaus (Iori and Kartalinsk) frequently occupied by mountain steppes, clay ephemeral deserts or semideserts, here called ‘‘steppes,’’ for example Shirak, 34 33 225) Mugansk, Adzhinaur, and others, and is confined particularly to the surrounding rugged foothills (bozdagi) with dry stream beds, erosion channels, and ravines. It also inhabits low, eroded clay mountains with desert or steppe vegetation, overgrown in some sections by juniper and pistachio; for example, the desert region of the Bozdag range. At places it ascends the mountains to heights of 1,800 to 2,000 and 2,100 т above msl (Kyal’vyaz in the Talyshin mountains; Dalidag near Istis, the Kel’badzharsk region of the Azerbaidzhan Soviet Socialist Republic; F.F. Aliev). In general, however, the hyaena avoids high mountains. It also does not live in dense forests, but is encountered in small groves and floodplain thickets on rivers, for example the Araks and Kur’, but rarely in gardens and vineyards. It has often been caught on the sea coast, for example at Vzmorsk station near Deshlagar and Derbent (Dinnik, 1914; Satunin, 1915; Aliev, 1971; Kh.M. Alekperov). In Turkmenia the hyaena lives along the stony desert foothills intersected by narrow valleys with sparse woody vegetation, arroyos, scarps, and along Fig. 7. Gorge in the Gyaz’-Gyadyk mountains (Turangovoe in the Kerlek area), habitat of hyaena and arkhar sheep (Ovis ammon cycloceros). Leopard and wolf also live here. Pistachio trees (Pistacia vera) grow along the slopes of the gorge. Badkhyz preserve, southern Turkmenia. May, 1962. Photograph by A.A. Sludskii. 35 26 rocky gorges almost devoid not only of woody and shrubby vegetation, but also grassy vegetation (Kopet-Dag, Gyaz’-Gyadyk, Badkhyz, Karabil’) (V.G. Heptner, 1956). It likewise occurs deep inside the mountains throughout the Kopet-Dag, but rarely and only at some places in the sandy desert of the Karakum, and more often in riparian thickets along rivers, for example among tamarisk at Kushka and tamarisk and poplar tugais at Tedzhen (V.G. Heptner). At Badkhyz, in southeast Turkmenia (about 850 т above msl), the hyaena inhabits the Er-oilan-duz and Namak-Saar basins and also Kyzyl- Dzhar plateau in the rolling countryside where high ridges alternate with broad open valleys, and very narrow and deep erosion channels. At places in these lowlands, weakly saline to saline springs are seen. Along the tops of the high ridges, and on their northern slopes, grow large, solitary trees of pistachio with crowns in the form of a dark green tent. Over the years the sandy loam soil of the ridges and valleys has become densely overgrown with a tall herbaceous carpet of meadow grass (Poa bulbosa), desert sedge (Carex pachystylis), and different varieties of sagebrush [Artemesia]. Gigantic fennels (Ferula badrakema) with peduncles as thick as a hand and reaching a height of up to 2.0 m are typical. In places, during years of good rainfall, they grow so densely that they impede the movement of man, and their thickets create the impression of a ‘‘forest’’. In some sections, above the low herbaceous cover, thickets of other large umbellifers grow, such as dorema (Dorema aitchisonii), the much smaller cousinia, bushes of Colligonum setosum, and black saxaul. This type of landscape is usually called a semisavanna.> Among vertebrates, reptiles of various species are found in large numbers in Badkhyz, in particular monitors and steppe tortoises. In some years both large and red-tailed gerbils are abundant, while thin-toed ground squirrels and sand hares are encountered occasionally. Even in the 1940s there were innumerable goitered gazelle, common mountain sheep [sic], and kulan. Numerous flocks of Karakul sheep are seen throughout the year. Hyaenas hunt some of these animals or feed on their carcasses. Large predators capable of competing with hyaenas, 1.е., wolf, cheetah, and leopard, are rarely seen. On Kyzyl-Dzhar plateau in May, 1962 hyaenas were sighted only once in 20 days of daily trekking, but the animal’s characteristic feces were detected several times. In Badkhyz, however, the hyaena also inhabits the low mountains of Gyaz’-Gyadyk, which are intersected by deep vaileys of small rivers with outcrops of limestone and sandstone. The steep slopes of these valleys are For photographs depicting the nature of the Badkhyz environment, also see sections on cheetah, leopard, and other species. 34 Ao Fig. 8. Rocky trail along bottom of a gorge used by hyaenas and also by porcupine, arkhar sheep, and other animals, sometimes even leopard. Kerlek region in Gyaz’- Gyadyk mountains, Badkhyz preserve, southern Turkmenia. February, 1965. Photograph by V.G. Heptner. covered with desert sedge and herbs. Along the gorges (down to the very bottom) of northern slopes grow centuries-old pistachio trees with trunks up to 1.0 m in diameter but a height of only 5.0 to 6.0 m; along the bottom of the gorges wild fig trees abound. Small saline springs occur in some gorges. In the larger gorges, for example in Kerlek, a narrow strip of reeds grow along the saline river bed. Hyaena tracks have been found at some of these springs as the animals use them as water holes and set up their dens in nearby rock fissures. The striped hyaena is also encountered in the deep basin (to 500 m) of Er-oilan-duz with a steep northern arroyo, and an undulating bottom relief. Low mud cones of greatly disintegrated volcanic rocks rise from it. The central portion of the basin is covered by extensive solonchaks. Along the edges of the basin grow small thickets of white saxaul, saltwort, and other plants. The grassy cover consists of desert sedge, sagebrush, and halophytes. During a week-long excursion, two fresh tracks of hyaenas were 36 35 28 detected in this basin and two old burrows and а den found under a projecting rock among mud volcanoes.+* In southwestern part of the Karabil’ upland, the hyaena inhabits extremely rugged, almost inaccessible, and unpopulated sites (Sheram- Kuyu, Shikhmulla, and Darvaza-kem wells, and many others), where flocks of Karakul’ sheep are driven for pasturing in autumn and winter (Nur- Gel’dyev, 1960). | In Turkmenia, however, the hyaena inhabits the extensive sandy desert, for example in the southeastern parts of the Karakum, where it is common in the black saxaul thickets at Lengych, Neder-Belent, Kert-Kuyu, Aitysh- Kuyu, and other wells at which it regularly or in some seasons snatches sheep (Sekunova et al., 1956; Nur-Gel’dyev, 1960). Thus on January 30, 1956, in the sands around the Lengych well in a trough 3.0 km x 3.0 km densely overgrown with black saxaul to a height of 3.5 to 4.0 m and thickness 30 to 40 cm, a pregnant female hyaena was caught. In this area hyaenas once lived rather permanently and were regularly seen from 1948 (Sekunova et al., 1956). Fig. 9. Lake Er-oilan-duz basin, habitat of hyaena. Arkhar sheep, kulan, and goitered gazelle are also found here. Badkhyz preserve, southern Turkmenia. May 1962. Photograph by Yu.K. Gorelov. ‘Data for Badkhyz are from V.G. Heptner, 1956 and unpublished material of A.A. Sludksii (1962) and V.G. Heptner (1942, 1948, and 1965). 29 In the extreme south of Uzbekistan and in southwest Tadzhikistan, the hyaena was confined roughly to the same biotopes as in Turkmenia. Moreover, in this region, it was noticed regularly in tugai thickets consisting of turanga, oleaster, tamarisk, giant plume grass, and reeds. Thus at the southern extremity of the Kugitangtau range, on the low mountain of Kulanashar, watered by the Amu-Darya, the hyaena was confined to tamarisk shrubs. Farther north this animal was found in the mountains at Kyzylalma village, in montane steppe with innumerable dryland and irrigated farms (Zarudnyi, 1917). In the southern spurs of the Gissar range, the hyaena inhabits the foothills of low mountains (Babatag, Aktau, Karshitau [Koktau], Garda-Niushti, and others) with steep forest-covered slopes and extremely rugged gorges and ravines, at the bottom of which saline springs are sometimes visible. The northern slopes are covered with steppe vegetation, and the southern with semidesert. There is a pistachio plantation in Babatag. The vertebrate fauna of these mountains is poor (Steppe tortoise, chukar and desert partridge, porcupine, mountain sheep), but [domestic] sheep are abundant (Arutyunov, 1964; A.A., Sludskii; A.K. Kovalev; A.I. Kirkhadzhi). In the tiger sanctuary ‘“Tigrovaya Balka’’ on the lower reaches of the Vakhsh, hyaenas regularly inhabit floodplains and different types of tugai vegetation. The highest elevations of the floodplains, sometimes with sand dunes, are overgrown with turanga trees. The trees in such a forest are sparse, seedlings and undergrowth are almost absent, and the herbaceous cover poor. In a turanga tugai one meets with small depressions and dry river beds, usually densely overgrown with tamarisk and some individual oleaster trees. Extensive depressions, periodically inundated by floodwaters, are densely covered with giant plume grass and more rarely by huge tamarisk bushes, which form almost impenetrable jungles. In the Vakhsh floodplain are many large fresh-water meanders and streams, the banks of which are densely overgrown with tugai or massive reeds and cattails. From the west, spurs of the Akatau range run almost up to the floodplain, and from the east, the Kashkakum sand desert. The vertebrate fauna in ‘“Tigrovaya Balka’’ sanctuary is very rich (monitor lizard, steppe tortoise, black-breasted pheasant, many ducks, sand hare, porcupine, coypu, goitered gazelle, and tugai deer). Many cattle and horses are here throughout the year. In the adjoining desert are steppe tortoise, gerbil, thin-toed ground squirrel, and other animals, likewise encountered in the floodplains, abound. In the sand along the floodplains [domestic] sheep are numerous (A.A. Sludksii). Two or three pairs of hyaenas regularly inhabit “‘Tigrovaya Balka’’ (46,000 hectares) (А.Т. Kirkhadzhi). Near the Pyandzh River hyaenas were encountered in a narrow valley between the end of the infertile Takakamar range and the river. Dense 30 tamarisk shrubs cover this gravelly soil impregnated with salt and at places filled with sand or broken rock (Zarudnyi, 1917). In Amu-Darya valley, in the region of Termez, this predator was formerly rather often encountered in reed beds and in ravines in adjacent river bank terraces (Sultanov, 1939). Thus throughout the Soviet Union the hyaena mainly inhabits foot- hills, clay ephemera deserts of the southern type, and montane steppes characterized by rugged topography and an abundance of dry river beds, gorges, precipitous scarps, and caves to serve as shelter. More rarely, the hyaena lives in sandy deserts, tugais, etc. The animal avoids high mountains and dense forests. Habitat selection largely depends on abundance and . accessibility of food—carcasses, fresh bones of wild and domestic ungulates, live ungulates, and steppe tortoises, and other animals. The hyaena, with its poor coat of sparse guard hair, thin underfur, and large ears almost devoid of hair, is little adapted to low atmospheric temperatures and cannot survive cold below —15 to —20°C. Hence it is distributed south of the January isotherm of 1.0°, does not extend high into the mountains, and inhabits only regions with a frostfree period of 230 to 270 days. The plains and low mountain regions of the Trans-Caucasus, Atreksk basin, the Kopet-Dag foothills, Tedzhen valley, the Murgab, upper part of the Amu-Darya valley and lower Pyandzh, Vakhsh, and several other rivers are prominent in the Soviet Union for length of frostfree period above freezing (Freikin, 1954; and others). The range of the hyaena within the Soviet Union is confined to these territories, and the animal seldom ventures out of them. Only on the west banks of the Caspian Sea, where the climate is mild because of the proximity of that water body, does the hyaena penetrate somewhat farther north. The absence of hyaenas in the northern half of the Karakum and the central and lower courses of the Amu-Darya is explained, apparently by prolonged freezes and short frostfree periods. To the north of the Karakum frosts of up to —20°С occur even in November, the temperature dropping sometimes to —25°C in January, and exceptionally even as low as —33°C. The mean January temperature is —2.0°С. The coldest weather occurs in the delta of the Amu-Darya where, at Nukus, the mean January temperature is —5.4°C. Within the range of the hyaena, at Termez the mean January temperature is 1.6°C and in extreme southern Turkmenia, even 4.7°C. The absence of permanent snow cover is also typical of hyaena habitats. In India, Pakistan, Afghanistan, Iran and Africa, the hyaena inhabits open desert expanses, especially in stony, rugged ravines. In Africa the animal is common even in the savannas. In forests it is restricted to edges or remains close to roads and paths (Pocock, 1941; Bourliere, 1955a; Prater, 1965). In Iran its vertical distribution extends up to 2,250 to 2,475 m above msl (Dinnik, 1914; Lay, 1967). 38 Si Food. Precise information on the food of the hyaena within the Soviet Union is scant. It is regarded everywhere as a carrion feeder, consuming mainly the carcasses of ungulates in different stages of decomposition, fresh bones, cartilage, ligaments, and bone marrow. The hyaena crushes long bones into fine particles and swallows them. Sometimes, though, whole bones or large pieces are found in its stomach. In the Trans-Caucasus, apart from carrion, it steals dogs, sheep, and other small domestic animals. Whether it attacks wild animals and birds in this region is not known (Dinnik, 1914; Satunin, 1915). In summer it consumes locusts and reptiles (Dal’, 1954). In a hyaena burrow in Badkhyz five horns of goitered gazelle were found, as well as a lower jaw and long bones, together with the shells of small steppe tortoises and the paws of red fox. Bones of a young wild boar were scattered in another four lairs and bones of mountain sheep in a fifth. In yet another hole the shriveled leg of a young kulan (onager) was found, and in still another the old bones of a mountain sheep were scattered. It is possible that the hyaenas did not catch these ungulates themselves, but merely picked up their bones (A.A. Sludskii). In one den the skull of a mountain sheep, bones of horses, and other bones were discovered. The stomach contents of one of two hyaenas caught contained the remains of a calf (evidently carrion), while the other contained a large porcupine quill broken in two (Yu.K. Gorelov). The stomach of a hyaena caught on January 30, 1956, in the southeastern part of the Karakum contained a pair of hooves, bones, and wool of a domestic sheep, and bits from the shell of a steppe tortoise. A male (in southwest Karakum) (of a pair, of which the female was caught) quickly bit to death two Karakul sheep (Sekunova et al., 1956). Similar information is available to show that in Turkmenia the hyaena feeds not only on carrion, but also hunts dogs, monitor and other lizards, and rodents, and consumes steppe tortoises, insects, and musk- and watermelons. In a single visit to a melon patch, this animal has been seen lopping off six or seven large watermelons (Heptner, 1956; Shcherbina, 1970). In southwest Tadzhikistan (Karshitau range), a young male caught on September 3, 1963 yielded stomach contents weighing 600 g, consisting of three large steppe tortoises, a small bit of skin with domestic goat wool, ten locusts, caper fruits, and a small amount of leaves and stalks (Arutyunov, 1964). The stomach of a young female caught on August 18, 1967 in the Babatag range contained the remains of 3 or 4 steppe tortoises, 18 locusts, and dry grain; the weight of this stomach was 300 g (АЛ. Kirkhadzhi). In the tugai along the Pyandzh, fecal examinations revealed drupes of oleaster (Elaeagnus angustifolia). The number of hyaenas increased in the period of ripening of these fruits in October-November in the tugais. These fruits are also consumed in large numbers by jackals, foxes, and jungle cats (Chernyshev, 1948). BZ In Iraq instances are known of this predator attacking horses and asses and also feeding on tortoises (Khett, 1959). In India the hyaena feeds mainly on carcasses of animals dead from diseases or killed by other predators; it also attacks dogs, sheep, goats, and calves (Jerdon,* 1874; Pocock, 1941; Prater, 1965). Many instances of hyaenas chasing leopards away from their quarry have been recorded (Pocock, 1941; Prater, 1965). In Africa carrion constitute the main food of the hyaena. It also feeds on large lizards, snakes, and fruits. It attacks, but not often, sheep, goats, and dogs (Ronnefeld, 1969). It is possible that the striped hyaena attacks live animals far more frequently than supposed. The animal’s nocturnal mode of life and innate caution make it difficult to assess the degree to which it is predatory, as is known in particular for the wolf in the Soviet Union. For example, it was long thought that the spotted hyaena (Crocuta crocuta) was a typical scavenger, but recent observations have established that it feeds mainly on carrion, only when living close to inhabited villages or in places where large predators are numerous. Elsewhere it actively and successfully attacks adult zebra, gnu, gazelle, or their young. In some regions of Africa hyaenas have destroyed up to 50% of the newborn antelope calves and older juveniles (Kruuk, 1966). Frequent finds of shells of steppe tortoises in hyaena stomachs in August—September and even in January when they are in summer—winter hibernation, suggest that the predators somehow detect the tortoise’s hibernacula and dig them out. Throughout most of the range of the striped hyaena, steppe tortoises are extremely abundant and constitute a ready prey for the carnivore. In southwest Tadzhikistan, after tortoises had gone into hibernation, one hyaena succeeded in capturing three after 2.5 hrs of nocturnal hunting around a water hole. Obviously, tortoises are an important constituent of the diet of the striped hyaena, especially when other sources of food are scarce. In zoological gardens the hyaena receives a daily ration of 3.0 kg of meat with bones (Balaev, 1940). Feeding mainly on carrion and bones, the hyaena evidently requires more water to quench its thirst than other carnivores. In southeast Turkmenia hyaenas drink from rivulets, springs, and wells in which water is close to the soil surface. In Badkhyz (southern Turkmenia) they are closely confined to saline springs; a liter of this water can contain up to 20 g of salts. The animals drink this water in the warm period of the year (A.A. Sludskii). Hyaenas inhabiting the southeastern parts of the Karakum are usually *Not in Literature Cited—Sci. Ed. 39 33 confined close to wells and drink water remaining in troughs after sheep have drunk (Sekunova et al., 1956). In southwest Tadzhikistan this carnivore drinks from rivulets, canals, and irrigation ditches; at places where fresh water is not available, it drinks from springs of saline water. It usually remains close to such springs in the warm season of the year. In winter and spring the animal drinks irregularly and, during this period, may be encountered 15 to 20 km away from the nearest water hole (Yu.K. Gorelov). Evidently, however, it will migrate long distances in search of water (Flerov, 1932). In the Kushka region of Turkmenia during the ripening of water- and muskmelons hyaenas regularly devour them, but they prefer the first, for which they enter the melon patch. Watermelon is mainly consumed for the purpose of quenching thirst (Heptner, 1956; Shcherbina, 1970). Water- and muskmelons were avidly consumed by a hyaena held in the Tashkent Zoological Garden (Balaev, 1940). Home range. The size of the breeding area during the reproductive period is not known. In search of prey a hyaena may wander far from its den. In ‘“Tigrovaya Balka’’ sanctuary in Tadzhikistan two to three pairs of hyaenas ranged over an area of 46,000 hectares. Outside of the breeding period, hyaenas are probably nomadic. However, in Turkmenia instances are known of great attachment by the hyaena to a given site. Thus, one hyaena lived for several years in a single lateral fissure of Gindovar gorge in the Kopet- Dag in the vicinity of Ashkhabad in a relatively populated locality. It is quite likely that such instances are due to the availability of convenient and safe hiding places, which by and large are few. The animal may be confined to a given place even after the mating season (V.G. Heptner). The hyaena marks its territory by leaving feces at dens, burrows, and in elevated places, such as the summits of hills and ridges along trails. Because of their high calcium content, the feces turn white rapidly and are visible from fairly long distances. Moreover, secretions from the special preanal scent glands of the hyaena also impart to the feces a characteristic odor. Such feces thus serve as scented and colored marks. Burrows and shelters. Throughout its home range, the hyaena establishes lairs in caves, rock fissures, and erosion channels, or in burrows formerly occupied by the porcupine or wolf, and also digs its own dens. Whatever previous animal may have tenanted an erstwhile hole, when a hyaena takes over occupancy, a characteristic collection of bones of wild and domestic animals will be found at the entrance (V.G. Heptner; Fig. 11). A burrow uncovered at Kurukulab (not far from Gurmab) in the Kopet-Dag was located in the central part of the broad valley of the Mergen’ul’ River. The entrance to the small cave, about 2.5 m high, was under a vertical wall of low rock. Farther in the cave narrowed and ran deep inside in the form of a tunnel. The lair was not visible from outside. At the entrance to the burrow and 42 34 among the rocks, many large bones of several species of animals were scattered in a semicircle 25 m in diameter; most were old and sun-bleached. They stood out sharply against the background of stones and the location of the burrow was visible some distance away. It was not visible from the road, however. The bones were mostly those of domestic animals (horses, cows and asses) but a few bones of mountain sheep were also present. The burrow had been used for several years (V.G. Heptner). In loose soil in ravines of the Kopet-Dag foothills, the hyaena sometimes digs its own burrow. The entrance is usually broad. In Turkmenia hyaenas often use the burrows of porcupines (V.G. Heptner). Six of the lairs examined in Badkhyz (southern Turkmenia) were identical to the one described above. One burrow was located on a completely exposed crest of a gentle ridge overgrown with low grass and solitary bushes. The hole had two entrances 60 cm x 40 cm. At both entrances there were large soil heaps measuring several cubic meters. Around the hole there were many feces in specially dug smal! pits. Scattered nearby were five pairs of horns of goitered gazelle, their long bones, tortoise shells, fox paws, and the skull of a porcupine. Another four lairs were found in broad fissures of limestone outcrops ina fairly deep gorge in the bed of the saline Kerlak River, on the northern slope of which grew old pistachio trees and at the bottom wild figs. At the entrance to one shelter lay the bones of mountain sheep and a juvenile wild boar. A lair found in the Er-oilan-duz basin was located on top of a small conical hill in a broad fissure among the outcrops of volcanic rocks. In addition to the main entrance this burrow had two exits among the rocks; around these lay the old bones of a mountain sheep and the legs of a young kulan. A few hundred meters from this lair, in a niche under a rock, was another hyaena den containing wool and bones of mountain sheep (A.A. Sludskii). At the same elevation hyaenas often occupied the holes of porcupines after widening them (Yu.K. Gorelov). In the southeast Karakum a hyaena burrow was found at the edge of a dense thicket of black saxaul. The entrance lay under a projection of compressed sand at the rim of an old well and was 67 cm wide. The hole ran to a depth of 3.0 m and extended in length for 4 m 15 cm. There were no chambers or enlargements inside. A second hole (2.0 km from Chaerla well) lay directly on the path leading to the well. The entrance to this was protected by low saltwort and saxaul thickets. Hyaena feces were evident around the hole and there were hairs at the entrance. The dimensions of the entrance were 72 cm x 38 cm. This burrow extended over 5.0 m in length and 2.5 m in depth (Sekunova et al., 1956). From time to time, in dense growth of bushes or reeds, the den for cubs will be open. In India the favorite sites for hyaena dens are caves among rocks or holes dug on slopes of hills or ravines. Most often hyaenas occupy and 40 Fig. 10. Burrow of a hyaena on а plateau in the Kyzyl-Dzhar area. Badkhyz preserve. May, 1962. Photograph by A.A. Sludskii. enlarge burrows of porcupines (Prater, 1965). In Africa hyaenas sometimes occupy burrows of aardvarks and wart hogs (Cotlow, 1960*). In the Tashkent Zoological Garden, just before parturition, a female began to dig a burrow with a male helping her in the task. Hyaenas dig the earth with the claws of the front paws and from time to time remove the dug-up soil with their hind legs. If small stones were encountered during digging, the male would pull them out with his teeth and drag them away from the surface. The hoie was dug quite rapidly to a depth of about 1.5 т; the female whelped in it (Balaev, 1940). During the day hyaenas hide in caves, niches, and holes, protecting themselves from enemies and the sun during summer and low temperatures in winter. Although nomadic, the hyaena rests during the day in temporary cover, such as ravines, niches, pits, dense thickets or reeds, and plume grass or Erianthus. In Badkhyz a hyaena was observed resting during the day оп an open piateau near a bush (A.A. Sludskii). In the hills around Kushka *Not in Literature Cited— Sci. Ed. 36 40 43 Fig. 11. Entrance to burrow of a hyaena on a level site (see Fig. 10). Around the burrow can be seen bones and horns of goitered gazelles. May, 1962. Photograph by A.A. Sludskii. during the day, hyaena sometimes hides under tumbleweed piles transported by the wind into hollows (V.G. Heptner). In the Karakum these carnivores rest among saxauls, hiding under wind-felled trees (Sekunova et al., 1956). In India hyaenas sometimes rest during the day among tall grass, shrubs, in sugar cane fields, caves, and under rocks (Pocock, 1941; Prater, 1965). Hyaenas often carry their catch to the burrow and consume it at leisure. Daily activity and behavior. The hyaena is essentially nocturnal and moves out from its den only after the onset of total darkness, usually returning to the shelter long before sunrise. It is rarely sighted during daylight hours. Once in Badkhyz a hyaena approached a hunter waiting in ambush during the day; the animal came so close that the hunter killed it with a blow to the head with his rifle (Yu.K. Gorelov). Hyaenas are cautious but extremely ‘‘cowardly.’’ Sometimes, however, especially during the night, they exhibit little fear of man. In Kushka instances are known when in twilight a hyaena will persistently trail a man along the road, keeping only a few tens of steps behind him. Hyaenas (at least in the 41 37 Fig. 12. Rocky gorge overgrown with pistachio, in which was located a hyaena breeding burrow. Kerlek region. Badkhyz preserve, southern Turkmenia. May, 1962. Photograph by A.A. Sludskii. 1940s) often came right up to houses in Morgunovsk village near Kushka, howled at their windows, etc. Hungry animals approached carrion very boldly (V.G. Heptner). In Badkhyz hyaenas formerly followed border patrols at night, within 20 to 50 steps behind them. It is possible that the animals were attracted to the viscera and heads of mountain sheep which the soldiers discarded from animals sometimes caught incidentally (Yu.K. Gorelov). In the event of unexpected danger the hyaena hides; it is very difficult to spot the animal in twilight and at night because of its striped coloration. The stripes crisscross the entire body surface, and therefore it is rendered inconspicuous. ‘‘Feigning death’’ is a characteristic pose of the striped hyaena. For example, if its lair is dug up and the hyaena exposed, the animal holds its breath, closes its eyes, and pretends to be dead; it can remain in that state for several minutes (Bourliere, 1955*). It also goes into this state when attacked by dogs from which it is unable to escape; in such a situation, *Not in Literature Cited— Sci. Ed. 41 Fig. 13. Rock (at lower right) under which lies the entrance to а hyaena den. Kamennoe gorge, Kerlek region. Badkhyz preserve, southern Turkmenia. May, 1962. Photograph by Yu.K. Gorelov. the hyaena suffers their powerful bites without resistance (Pocock, 1941). When the dogs finally leave, it jumps up and runs away. A tamed hyaena fondled by its master and sexually aroused everts its rectum to a length up to 5.0 cm. Evidently, such behavior is a form of communication between these animals since special scent glands producing secretions are located at the anal opening. During rut hyaenas evert the rectum and press it against prominent objects, thus marking their own territory, within which they remain. When an enemy attacks, the hyaena similarly everts the rectum and sprays secretions from the rectal glands, which give off a strong unpleasant odor (Ronnefeld, 1969). While seeking quarry, the hyaena is mainly guided by its olfactory sense which, however, is not as well developed as in dogs. Vision and hearing are also good although some researchers report that hearing is relatively weak (Pocock, 1941). Hyaenas feeding on the remains of prey captured by other carnivores, often remain in their vicinity, forming a sort of commensal relationship. Such groups of hyaenas are common in places in southern Turkmenia where wolves are quite numerous. 42 39) Fig. 14. Bones at entrance to a hyaena burrow (see Fig. 15): horses, goitered gazelle, arkhar sheep, wild boar, and other animals. May, 1962. Photograph by Yu.K. Gorelov. There is no information about the methods hyaenas employ for hunting live prey in the Soviet Union. In other countries they usually hunt in pairs or family groups of five or six animals. They are fleet-footed and hardy. Having reached the prey by chasing, the predator grabs it by its flanks, in the region of the groin, inflicts large mortal wounds, and pulls out the viscera, which causes the victim’s death (Krumbiegel, 1954). Hyaenas live in pairs, more rarely singly, or form small family groups of five or six animals. The pair lives in a single den. In captivity hyaenas often quarrel among themselves. One method of attack is to attempt to hold the opponent by the leg. In this process the attacker approaches sideways. Another method is the throat hold. During fights hyaenas often inflict severe wounds on each other, but then upon calming down, rest peacefully in one den (Dalaev, 1940). Hyaenas caught when young, are easily tamed and behave like dogs even after attaining adulthood. They are amenable to training. Seasonal migration and transgressions. These aspects are not known. In southeast Turkmenia, at Badkhyz, following the drying up of water sources, fires, and the migration of wild and domestic ungulates to the 44 40 Fig. 15. Rocks among which а hyaena den was found. Er-oilan-duz basin, Badkhyz preserve, southern Turkmenia. May 1962. Photograph by A.A. Sludskii. Kushka valley, hyaenas and wolves follow the migrants. Thus in the second half of July, 1942 at Morgunovsk village, hyaenas appeared in larger numbers than previously. They were sighted very often in tamarisk thickets. These carnivores arrived there from the desert. Such a migration of hyaenas to this river was not seen in 1941 (Heptner, 1956). The hyaena population increased during the autumn in the tugais of the Vakhsh, Pyandzh, and other river valleys in southwest Tadzhikistan. This phenomenon may be explained by their migration from the surrounding desert. Hyaenas also congregate in areas of heavy mortality of wild and domestic ungulates during outbreaks of epizootic diseases or during heavy sleet. In Africa, together with predatory animals such as lions, cheetah, and hunting dogs, hyaenas accompany herds of migrating ungulates, attack sick and weakened animals, and subsist on the carcasses of the dead. Reproduction. The striped hyaena is monogamous. The male helps the female in establishing the den and in raising the young. The periods of rut reported in literature are only partly true and require verification, viz., November—December and January —February for hyaena in Trans-Caucasus (Ognev, 1931; Aliev, 1971) and October-November in southeast Turkmenia (Shcherbina, 1970). According to observations made _ 45 41 in the Tashkent Zoological Garden, periods of rut in the striped hyaena are not seasonal. A female mated three times in 1937: January 11, April 15 (three days after the death of her litter), and December 16. She whelped on all the three occasions. A female mated twice in 1935: in the middle of June and on December 30 (Balaev, 1940). These observations suggest that, under natural conditions, hyaena young may be seen in all seasons, which is typical of many carnivores in the subtropics and tropics. Coitus lasts for over an hour. During mating the male grips the skin of the neck of the female in his teeth; it is not restricted to any particular time of the day. During the period of mating the animals remain irritable. Gestation requires 90 to 91 days (Heck and Hiltzheymer, 1925; Balaev, 1940; Krumbiegel, 1954). In the Tashkent Zoological Garden a female mated on January 22 and whelped on March 16. In the Trans-Caucasus cubs appear in April-May (Ognev, 1931). In May a pregnant female with four well-developed fetuses was killed there (Satunin, 1915). In Turkmenia, in the southeastern part of Karakum, on January 30, 1956, a large pregnant female with considerable subcutaneous and intestinal fat was killed. She carried three fetuses 9.8 cm in length and 1.0 to 1.5 - months of age (mating had evidently taken place in December; Sekunova et al., 1956). In southeast Turkmenia pregnant females were caught in January and lactating ones in February (Shcherbina, 1970). In southwest Turkmenia hyaena litters were seen in March (G.P. Dement’ev, 1955). The litter invariably consists of one to four cubs, usually three. In the Trans-Caucasus two to four cubs are common (Satunin, 1915); in southwest Turkmenia two or three, and in the southeast two to four (Shcherbina, 1970). In the Kopet-Dag a litter contained two cubs (V.G. Heptner). In the tugais of the Vakhsh valley (southwest Turkmenia), a lair with four cubs aged 1.0 to 1.5 months was found among reeds in early June, 1950 (Chernyshev, 1958). In southern Uzbekistan hyaena litters consist of three or four cubs (Sultanov, 1939). According to observations in the Tashkent Zoological Garden, the hyaena produces one to four cubs, but as a rule three. Thus one female in her first whelping birthed a single cub, in the second, four, and subsequently three each time (Balaev, 1940). In India hyaena litters consist of two to four cubs (Blanford, 1888; Pocock, 1941) but three or four in different parts of Africa (Ronnefeld, 1969). Whelping lasts, on average, for two hours, commencing usually between 10:00 a.m. and 12:00 noon. Growth, development, and molt. Young are born blind, with closed ear passages, and are the size of the pups of an average-sized dog. The hair coat of the newborn is whitish-gray with distinct black stripes. Exposed to the light, cubs immediately whine like pups and constantly crawl around the den. They begin to see clearly on the seventh or eighth day. By the tenth 46 42 day their measurements are as follows: female No. 1—body length 30 cm and tail length 7.0 cm; female No. 2—29 and 8.0 cm; and one male 29 and 8.0 cm respectively. Up to the age of one month the young are confined to the den and only later begin to venture out. At first they are very cautious and hide in the den at the slightest sound. They emerge from it singly, mainly on the leeward side, and before leaving peep out to examine the surrounding area. In sheltered localities in the Kopet-Dag, growing cubs often play around the burrow during the day (V.G. Heptner). The young, up to the age of two months, feed exclusively on their mother’s milk and begin to take meat only thereafter. The prolonged period of lactation is an adaptation since cubs, in their early life, cannot feed on carrion, especially bones. Both parents participate in feeding the young, by the female more so and for a longer period. Before the young emerge from the den the female remains with them continuously; if she leaves them at all, it is only for a very short while. She is fed during this period by the male. In the presence of the female the male will not approach the cubs since she invariably drives him away. In her absence, however, the male sneaks to the cubs, sniffs them, and licks them. When the cubs begin to emerge from the den, the female follows them vigilantly. She does not allow them to remain outside for long and, picking them up by their neck with her teeth, pushes them back inside the den. In spite of the male’s solicitous attitude toward the cubs, the female is suspicious of his intentions and invariably chases him away whenever he approaches her litter. The female eats the feces of her cubs (Balaev, 1940). By autumn young hyaenas are roughly one-half the size of their parents. Thus a young male caught in southwest Tadzhikistan on September 3, 1956, aged about 5 months, had the following measurements: body length 94 cm, tail 38 cm, ears 11 cm, and weight 19.4 kg. Its deciduous teeth had already been replaced by permanent ones and the premolars lost (Arutyunov, 1964). A young female caught in the same region on August 18, 1967 had a body length of 91 cm, tail 36 cm, ears 11.7 cm, and weight 17.2 kg (АА. Kirkhadzhi). In India an adult male of 150 cm body length weighed 38.5 kg, and a female 34 kg (Prater, 1965). The period of the onset of sexual maturation in the hyaena has not been accurately established. In the Tashkent Zoological Garden a female born in 1929 produced her first litter in 1933, 1.е., in her fourth year (Balaev, 1941). Hyaenas live for an average of 12 years but have survived for 23 in a zoological garden (Flower, 1931). Molt has not been studied. Enemies, competitors, diseases, parasites, mortality, and population dynamics. Within the Soviet Union wolves are evidently enemies of the 47 43 hyaena and, until recently, so were tigers and leopards. The main competitors for their food are wolves, jackals, red foxes, rarely corsac foxes, and griffon vultures. The importance of griffon vultures where they are abundant, for example in Badkhyz (in the 1940s), is paramount. Carrion remaining at dawn is consumed by vultures in the early hours of the morning, leaving no remnants except bones. The following example reveals the competitive relationship between hyaenas and large carrion-eating birds, primarily griffons and other vultures. In Badkhyz preserve in the early 1940s, when there were large numbers of these birds, hyaenas were also common. Goitered gazelle served as the major food source for all. After the 1950s and early 1960s goitered gazelle became almost extinct in southern Turkmenia and by 1965 nesting colonies of griffons had decreased sharply and hyaenas had become almost extinct (V.G. Heptner). At the same time, as previously mentioned, hyaenas often become the commensals of tigers, leopard, and cheetah, since they finish off the remains of carrion, especially bones, left behind by these predators. While wolves, like hyaenas, crush most bones and consume them, large cats do not. Diseases and parasites are almost unstudied. It is known only that hyaenas are highly resistant to various infectious diseases and suffer only rarely even when held in zoological gardens (Balaev, 1940). A hyaena caught on January 30 in the southeast Karakum was infected by Taenia pisiformis in the abdominal cavity; fleas (Synosternum pallidus) were also collected from the same animal. The substratum collected from the den of a hyaena in the same region at a depth of 3.0 m revealed the presence of mites (Eulaelaps stadularis and Haemalaelaps longipes) and ixodid ticks (Hyalomma а. asiaticum) (Sekunova et al., 1956). From an animal killed in Badkhyz on January 23, 1960, two Ixodes ticks and five Pulex irritans fleas were removed (Yu.K. Gorelov). In severe winters with plentiful snow, when heavy sleet affects even wild and domestic ungulates, the warmth-loving hyaena is not adapted to deep snow cover and perishes. At places where hyaenas live near water holes, they sometimes die if buried under ice. For example, in the last century in the Trans-Caucasus they ‘‘оНеп drowned in Lake Karayaz by falling through the ice; three bodies of dead hyaenas were found one spring’’ (Ognev, 1931). Population dynamics are not clearly understood. Field characteristics. The size of a hyaena is that of a large species of dog, for example the German shepherd. The head is massive with broad and blunt muzzle and large triangular ears pointed upward. Along the middorsal neck and back there is a well-developed, erect mane. The anterior part of the body is strongly elevated and the rear set low, as a result of which the animal seems to rest on its hind legs. The front legs are bent. The short 46 44 tail is equal in length to the thigh and shaggy. From а distance the hyaena appears almost white; dark transverse stripes on the flanks and the erect mane on the neck are only visible in good light. The animal walks or trots. When frightened, it runs in a characteristic heavy gallop, tucking its tail between the hind legs, turning its ears back, and exudes a strong unpleasant odor. The footprints of the forefeet are considerably larger than the hind ones. Undisturbed footprints of the forefeet in a solonchak measure 10 cm in length and 8.0 cm in width; corresponding measurements of the hind feet were 8.0 and 6.0 cm. When the animal is walking quietly, the footprints of Fig. 16. Feces of hyaena with characteristic white coloration. Plateau in Kyzyl- Dzhar area (see Figs. 10 and 11), Badkhyz preserve. May, 1962. Photograph by A.A. Sludskii. 45 the hind feet do not coincide with those of the forefeet (as in wolves, jackals, and foxes), falling somewhat behind or sideways. At a normal pace the distance between imprints of front and hind feet is 55 to 63 cm. Usually, impressions of the four claws are visible in all footprints. Feces of hyaenas that are feeding on bones are very typical (Fig. 16). They are white, in the form of slightly flattened balls, varying in diameter from 0.5 to 2.5 cm, and usually consist almost exclusively of disintegrated bones. They occur in heaps and are often found in pits around the burrow (A.A. Sludskii). The voice is a typical cracked howl which can be heard, though rarely, at night in all seasons of the year. Many hunters cannot distinguish it. A wounded animal growls. (A.S.). Practical Significance The pelage of the striped hyaena is coarse and sparse; it is not considered a fur-bearing animal. Hunting for hyaena is not specially organized; the animal is incidentally caught in traps set for other animals. It also dies of poisons used in baits for wolves, or is chased and shot. It is caught from time to time with the help of hunting dogs and killed later, at a more convenient time. As a matter of fact, the hyaena has no fur and hence the few skins sold by hunters are generally thought to be wolf or dog skins of poor quality. Depending on the quality of the skin and its defects, the selling price ranges from 45 kopecks to 1.0 ruble 80 kopecks. Hyaena skins are used for preparing chamois leather. Many hyaenas were formerly caught in Turkmenia, and about 130 skins were collected between 1931 and 1937. Commencing from 1948 in this republic, only a few tens of skins were received for curing, and almost none received by 1970. In the Trans-Caucasus, Tadzhikistan, and Uzbekistan, only a few stray hyaenas were caught in the 1960s and their skins seldom sent for curing. For the Soviet Union as a whole, in the 1930s about 200 skins were processed, less than 100 in the 1950s, and processing actually ceased by 1970. This animal is of considerable interest to zoological gardens. Live hyaenas are mainly caught in Turkmenia and in small numbers in southwest Tadzhikistan. Five hyaenas were caught in Tadzhikistan in 1949 and 1950. In Turkmenia female hyaenas were earlier highly prized. A belt cut from the abdomen together with the genital region, conforming to certain patterns, was considered a powerful talisman and was in great demand. Magical properties were attributed even to the tail. Dead hyaenas were thrown away but the tail always removed (V.G. Нершег). The hyaena is little known as an enemy of animal and game husbandry 48 46 in the USSR. In Turkmenia the animal causes damage insofar as it steals foxes, cats, hares, and even goitered gazelles from traps, making off with the trap (V.G. Heptner). At present, because the hyaena population is small over much of the range, the damage these animals cause is negligible. In India hyaenas living in the environs of villages cause ‘“considerable damage’’ by destroying goats, sheep, calves, and small dogs (Pocock, 1941); they are thus a “‘real enemy of livestock.’’ The damage they inflict on melon patches by feeding on water- and muskmelons is insignificant. In the last century the hyaena had a very bad reputation as a man- eater. In the 1880s the newspaper “‘Kavkaz’’ reported that some predatory animal had attacked human beings, especially sleeping children, for three years in Igdyr’ Erivansk province; in just one year 25 children were wounded. Three adults were also bitten in the same region. Attacks invariably occurred at night on people sleeping in the open, in gardens or farms. The hyaena was labeled the culprit. The police administration announced a reward of about 100 rubles for every hyaena killed. Attacks on man by hyaena were reported later from other areas of the Trans-Caucasus, e.g., Surmalinsk district of Erivansk province; such attacks were also reported in 1908 (‘‘Okhotnich’ya Gazeta,’’ nos. 25 and 38). Those well acquainted with the fauna of Trans-Caucasus (Dinnik, 1914; Satunin, 1915) considered the foregoing information reliable and also blamed hyaenas for these deaths. Instances of hyaena stealing and killing children sleeping in courtyards were reported in Azerbaidzhan even in the 1930s and 1940s (Piraze Agdash region, 1939; Mardzhanly Dzhabrail’sk region, 1949). In 1942 in the Ismailinsk region (Golyndzhakh) a hyaena entered a hut where the guard was sleeping and mauled him (F.F. Aliev). In southeast Turkmenia (Badkhyz) hyaenas have reportedly carried away small children during the night (Yu. K. Gorelov). The carrying away of a child by a hyaena around Serakhs in Turkmenia was reported as late as 1948 (V.G. Heptner). Instances of stealing of small children have also been reported in India (Pocock, 1941). Hyaena attacks on sleeping children in Tadzhikistan and Uzbekistan have not been confirmed. Sometimes wounded animals attack man (Dinnik, 1914; Satunin, 1915). Hyaenas sometimes dig up and eat human bodies when the burial is shallow and no wooden coffin is used (Berkhnii Guzlak, Fizulinsk region, 1936; Lembran, Bardinsk region, Azerbaidzhan; F.F. Aliev). In ancient Egypt striped hyaenas were domesticated. They were raised as livestock on special food. The habit of feeding hyaenas continued in North Africa up to the 20th century and was practiced, for example, by the Tuaregs (Ronnefeld, 1969). At present, over much of the range in the USSR (Trans-Caucasus, southwest Turkmenia, Tadzhikistan, and Uzbekistan), the hyaena has become 49 47 а rare animal and causes almost по harm to the national economy. It poses very little danger now even for human beings. As a rare and extremely interesting animal to scientists, extinction of the hyaena should be prevented throughout its range in the Soviet Union. In Azerbaidzhan Soviet Socialist Republic the law (1970) bans their hunting year-round and a severe fine is imposed for unauthorized killing. Catching a hyaena when it attacks man or domestic animals and licensed shooting for scientific purposes are permitted. (A.S.) Family FELIDAE Gray, 1821 Felidae are highly specialized carnivores, in fact the most specialized family in the suborder and order. Size is small, moderate, or large. The smailest species has a body length of about 50 cm and weighs 2.0 to 3.0 kg, while the largest species (Amur tiger, Panthera tigris altaica) measures up to 300 cm in length and weighs up to 390 kg (ratio of body length between the smallest and the largest 1:6 and weight 1: 130 to 195). The dimensions of the tiger are inferior only to those of brown or polar bears. All species of the family, even those widely differing in external appearance (lion and cheetah) have an extremely characteristic, typical feline appearance, and are essentially monotypic. With a fairly large number of species (35), this family is incomparably more uniform than the family with the greatest species richness—viverrids (Viverridae, about 75 species), or weasels (Mustelidae, about 70 species) and even than dogs (Canidae), comprising fewer species (29). Felids are of relatively light build, very agile, with elongated (in some cases very long) trunks, and usually a long tail. Tail length is not less than one- third, and sometimes more than one-half, the body length, and less than one- third only exceptionally. The legs are not long, but powerful, and the general outline of the body can be described as a parallelogram, sometimes extended. In some species (lynx, caracal, and especially cheetah) the legs are long and the body profile squarish or nearly squarish. The thorax is small in volume and not prominent (lesser volume of lungs and heart); the body appears laterally compressed and the thorax is no larger than the abdomen in volume. The forelegs are straight while the hind legs in most species are somewhat more powerful, although the angles between the foot, shank, thigh, and pelvis are relatively small. The sacral region is raised and stands above the shoulder region; the posterior portion of the body appears particularly powerful, more powerful than the anterior. The dorsal profile of these species (a majority) is elevated posteriorly, being sometimes (quite rarely) almost straight. Such a particularly full development of the animal body favors a powerful leap from a standing position (without running), 50 48 which is typical of all smaller species and some larger ones (рита, snow leopard). In a few species (big cats, i.e., lion, tiger, leopard,! and jaguar), the anterior region is more powerful, the back slopes upward anteriorly, and the shoulder region stands above the sacral, or more rarely the dorsal profile is straight (Fig. 30). Neck of moderate length or short. Head relatively small, with a shen facial portion, and muzzle short and blunt. In some forms (some cats of genus Panthera) facial portion relatively large, elongated, and massive, but not so extended as in members of the wolf family. In many, the facial portion is so shortened that the shape of the head is almost spherical. In all these cases the chin region is shifted to the level of the profile of the bare nasal region; this is due to the position of the incisors and the position and intense development of the canines. ' Eyes large; much larger and protuberant in smaller species; pupil rounded or slit. Ears highly variable in size and form; in some (large species) they are not very large, sometimes almost concealed by hair (lion, tiger, and snow leopard in winter), but are usually (small species) large, rounded or pointed at the tip, set at a slight angle to the sagittal plane, or more rarely very highly inclined sideways (“‘hung out’’). The ears, when of a significant size, are sometimes set erect, or almost so, pointed, and carry long hair at the tip (tuft as in lynx and caracal). In some the ears are set erect, rounded, and very large (several species). The total area of the ear opening may be more than the area of forward projection of the head. From the outside the pinna is slit in the middle and the antitragus well developed, simple, and with a cylindrical thickening. The considerable development of the external ear corresponds to a significant development of the bony resonators; hence the sense of hearing is one of the most important senses of various species of the family. Lip (‘‘whisker’’) and brow vibrissae are well developed, particularly the former, oy in small species; mandibular vibrissae are absent. Limbs digitigrade. Forelegs with five digits each, one of which is set high and very small. Hind legs with four digits each. Paws of fore- and hind legs (especially the former) powerfully developed, digits wide-set (paw spread out and not compact as in wolves); and digital pads and large triangular sole pad well developed. Single small carpal callosity present; metatarsal callosities absent. Lower surface of paw usually hairless, and callosity in most species well formed; its impression is well seen in pug Throughout this section the term bars [leopard] is used, which has been reserved from early times for the species Panthera pardus. Recently, it has often been written “‘leopard”’ in the Russian language, a foreign word alien to the Russian tongue. Nowadays bars is being employed for Uncia uncia. For this species I have used the term snezhnyi bars [snow leopard] ог irbis [ounce] (see р. 276). 49 marks. However, in some species, especially in winter (lynx) ог even year- round (sand cat, Felis margarita), hair on lower surface of paw so dense and long that it completely conceals the pad. All four digits on each limb roughly similar in size; digits III and IV longer but equal in length, and Папа У equal in size but somewhat shorter than middle ones. On the whole cat paws are broad and rounded. All the digits are interconnected with an elastic dermal membrane developed to different degrees but, in the majority of species, reaching digital pads or claws (see Fig. 278). Digits are extremely mobile and can move widely sideways. This is an important feature of the cat’s paw as a grasping organ. Cat digits bear powerful claws. They are relatively short but strongly curved, sharply compressed from the sides, and very sharp. The claws in all Russian cats, except cheetah, are retracted; i.e., in the resting position, and during movement they are sharply drawn backward and raised. Their sharp tips, though pointed generally downward, do not reach the ground. Cats move on the pads present at the joint between phalanges [ and II. The special articular surfaces of the phalanges permit a very broad independent vertical movement, while a system of powerful ligament permits retraction and extension of their claws (Fig. 17). Two large ligaments extend along the top and bottom of all the phalanges, which are firmly attached in front to phalanx |; these, in addition to a system of tiny ligaments connecting phalanges I and II, play a special role. In the resting position phalanx III is set at an angle of about 45° with its distal end turned upward; phalanx П in relation to Ш forms almost a right angle and is set downward. Phalanx I is pressed closely to phalanx II. The main axis of the former is parallel to that of the latter and raised up and slightly back. In such a position the claw is retracted. When the claws are extended (a volitional act) by activation = WZ . 7 Pa Fig. 17. Skeleton and ligament of digits of tiger showing position and mechanism of claw release. Upper sketch shows position at rest with claw retracted; lower sketch shows claw extended. Anatomical data from V. Mazak, 1965. Sketch by N.N. Kondakov. all 50 of the ligaments, mainly the lower one, phalanges III and II align; phalanx I, in relation to II, is so positioned that it is vertical and somewhat projected forward. The claw is extended but its tip bent downward. The phalanges as a group can also bend downward. The horizontal flexibility of the phalanges, extension of claws, and downward bend of the digits make a cat’s paw a powerful and complete grasping device. The clasping capacity of a cat’s paw is mostly determined by the form of its claws. Their curvature varies but is always strong, in several the outer margin of the claw forming a smooth arc of small radius. In several species, especially in large cats, the claw is firmly attached to the phalanx in such a way that its base is enclosed in a well-developed, small bony collar. On the whole, the gripping mechanism is better developed in the front paws, which also play the role of chief grasping instruments (Fig. 17). In a typically well-developed cat’s paw each claw has a well-defined, highly extensible skin fold which covers the retracted claw laterally and dorsally. These ‘‘sheaths’’ protect the claw and its tip. Their ‘‘stretch’’ when the claw is released is quite remarkable.? In species in which such sheaths are well developed the interdigital membranes are likewise generally well developed and reach the digital pads and claws. In some species, however, claw sheaths are wholly or partly reduced, occasionally even poorly developed. Interdigital membranes may also be reduced to some extent. Both structures are greatly reduced in cheetah (Fig. 278). The paw of this animal is characterized by other features—claws less sunken and externally visible (see characteristics of cheetah). Claws come into play only during gripping, holding, or killing prey, or in self-defense, and to a lesser extent when climbing trees; they are not used ai all in locomotion or for digging. Thus their tips are always sharp and ready for action. At the same time, the anatomy of the paw is probably one reason why cats cannot run fast or long. Only cheetah are fleet-footed and hunt prey by sprinting. Among mammals, they are the swiftest animals. A paw with retractile claws is a characteristic feature of the family; among other carnivores, this feature is seen only in a small group of viverrids (Prionodontinae; genera Prionodon and Pardictis). In most animals the pelage is dense, soft and silky. The length, compactness, and extent of growth of underfur vary considerably, depending on climatic conditions in the habitat. In severe climates seasonal differences in coat color are distinct. The length of hair throughout the body is fairly uniform in most species. Only in a few (lynx) are long ‘‘side-whiskers’”’ found on the cheeks. In male lions a dense, long mane occurs on the anterior "Some references in literature to the presence of horny caps at the base of claws (Sapozhenkov, 1957; and others) are incorrect. They are evidently mistaken descriptions of the dermal folds described above. Sil part of the trunk, neck, and head up to the ears; long hair also occurs along the chest and abdomen. In cheetah elongated hair likewise occurs along the nape and on the withers. Coloring extremely variable, but mostly spotted, with dark spots lying on a fundamentally light background. The shape of the spots varies greatly, from simple, close-set, small dots, to large ring-shaped areas consisting of individual spots, completely circular or elongated rings, and so on. Quite often the spots form fairly regular longitudinal or transverse rows; some merge into transverse or longitudinal stripes; coloration is thus a combination of strips and stretched and circular spots. The trunk region of some species is devoid of spots, consisting only of vivid transverse stripes (tiger). The general background color is usually a light shade, varying from almost white or light yellow to rusty, bright rusty, or cinnamon. Spots are usually black, of varying intensity and sharpness of outline, and sometimes rust-tinged. Some species are uniform in color (caracal, puma and lion) or almost so (sand cat, Felis margarita); their color is usually sand-yellow, brownish-yellow, or some similar shade. Sometimes against a uniform general background very poorly developed spottedness is visible, or only traces of it on the limbs (some forms of lynx), or a pattern on the head (one of the colored forms of Temminck’s cat, Felis temminckit). Usually, even among monochromatic species kittens or juveniles have more or less developed spots. The spotted pattern of present-day cats should be considered as an ancestral character. Individual color variability is pronounced in many species, as is geographic variability. Geographic variability is generally exhibited in a change in overall color shade, and often in type of spottedness. In some species melanism, albinism, and chromism are encountered, while geographic ~ localization of such abnormalities is known [black “‘panthers’’ (leopards) of India, chromic manuls of Turkmenia, and others]. Some species show seasonal variation in color (Felis yaguarundi of America). In one case (Felis temminckii) the color, although not being in the strict sense polymorphic changes very sharply (spotted to uniform, dark to light); geographic localization of color types is also known. Among lions sexual dimorphism is manifested in the growth of a mane in males. In all cats males are larger than females. Anal glands are usually present but poorly developed. Penis located directly in front of the scrotum. Glans rudimentary and with horny spines. Teats, two to four pairs. The skull is characterized by several distinct family features. However, as a result of significant differences in the absolute overall size and other characteristics of different species, the general appearance and skull proportions may vary considerably. In general features the skull is broad, 52 with widely diverging zygomatic arches. Anterior part of skull, from anterior margin of orbit (‘‘muzzle’’), short and broad. Ethmoturbinal scrolls large, extend forward almost up to opening of nasal chamber, and lie on top of maxilloturbinals. Alisphenoid canal absent. Bony auditory bulla usually high, rounded (swollen), formed of ectotympanicum and entotympanicum, and divided internally into two chambers by a septum. Entotympanic (rear-medial) portion invariably, but to varying degrees, larger than ectotympanic portion, which communicates with ear opening. External separation of two portions of bulla not usually perceptible and distinct only in some forms. Auditory tube absent or barely discernible. Relative size of tympanic bulla highly variable; large, swollen, and thin-walled in small species (especially in desert species); relatively small in large animals and often flat and thick-walled. Mastoid and paroccipital processes separated; latter flat, usually closely adhering to posterior wall of bulla, and in a majority of species does not project beyond lower margin of tympanic bulla. The following are general changes in the overall appearance of the skull. Skull of more or less neutral type (lynx, caracal; Vereshchagin, 1967): general outlines fairly elongated—oval; cross section about 75% of condylobasal length. Interorbital and postorbital constrictions broad, cranium very voluminous, weakly extended forward, and highly enlarged; zygomatic arches widest at level of articular surfaces but their general outline bulges or is arched. Crests and tuberosities for attachment of muscles, especially masticatory ones, weakly developed; sagittal crest present but small, while lambdoidal crest well developed. The series comprising species of larger and largest cats and culminating in tiger is characterized by the skull being very heavy and massive, the facial region, especially the muzzle, relatively elongated, and the cranial region relatively short. The entire skull assumes a highly extended form although the rear portions of the zygomatic arches may be very broad. Viewed from the front the features are not so bulging but appear very angular. The interorbital and especially the postorbital constrictions become relatively small and volume of the brain case is reduced and not inflated. Tuberosities and crests very powerfully developed; sagittal and lambdoidal crests particularly large and high; brain case greatly compressed by powerful masticatory muscles. Teeth, especially canines, powerful with a broad base, but relatively short. Tympanic bullae relatively small, flattened, and thick- walled in extreme cases. Lower jaw heavy, relatively long, with a powerful anterior portion projecting forward, bearing large canines with long and massive bases. This is a skull typical of animals which kill and tear apart large quarry. Often the quarry is larger than the predator in size and weight; in the very largest by several times. 53 53 The skull of small species is characterized, on the contrary, by lightness, a thin-walled brain case, and absence of any crests or tuberosities, apart from the lambdoidal. Sagittal crest not developed, or represented only by a small ridge at occipital end and lyrate figure [temporal crest] usually broad. Facial region of skull abbreviated to a greater degree, muzzle short and blunt, and orbits highly enlarged and increasingly set forward. Zygomatic arches very thin in midportion (not only from the rear) and form a fairly regular arcuate shape. What is important is that the brain case, sometimes moderately enlarged and slightly extended, becomes increasingly voluminous, bulging, and short. Thus the entire skull is short and swollen with broad interorbital and postorbital regions (see Fig. 19). The general contour of the skull in some forms is roughly oval, as in lynx, or even somewhat elongated (elongated brain case), or tends to become broader and shorter, and in extreme cases (manul, F’. manu!) even spherical. On the whole the skull of small cats, compared with that of large ones, appears infantile. The teeth are relatively weak and the canines relatively long, but thinner. The lower jaw is short and light, with a relatively weak, not massive, anterior portion. This is the line of specialization of small predators which kill small prey, much smaller in size (rodents) than the predator itself. If cats are regarded as perfect predators the above two forms are superior types, even though morphologically they contrast with each other. Dentition shows extreme specialization within the order with regard to number of teeth and certain features of their structure. Complete dental formula: 1 3 m m _1 he ne aay, ee — 30, 3 but in some cases second upper premolar (first in row since first upper premolar generally absent in cats) sometimes does not grow (or is lost), or is invariably absent (manul and lynx). Dentition in these cases: п и 3 1 2 1 The diastema, especially in the lower jaw, is large in most species. This is associated with the absence of the first premolar in the upper jaw and the first and second premolar in the lower jaw, which in turn is due to the intense development of canines and jaw abbreviation. The teeth are powerful and sectorial (cutting). Carnassial teeth large, highly compressed, and relatively long; incisors small and weak, form straight transverse rows, and canines large. In large species they are more 55 54 massive and short and in small species relatively thin and long, sometimes very long and thin (clouded leopard), and usually with shallow longitudinal grooves. Second premolar (upper) very small, conical and single-rooted; third upper premolar well developed, compressed from the sides, usually with a high and sharp central cusp, and two small cusps located at its base at front and rear. Upper carnassial tooth (fourth premolar) very large, compressed from the sides, with three cusps lying in a single plane; first cusp moderate and conical; second high and sharp; and third low and long with a cutting edge. Antero-internal surface of teeth, in most, with additional cusp with a small, usually well-developed blunt cusp. First upper molar rudimentary, very small, and set across the row (sometimes lost). In the lower jaw both premolars (third and fourth) have three cusps, like the third premolar in the upper jaw. Carnassial tooth (first molar) powerful with two well-developed cutting edges. Milk carnassial in some species complex compared with permanent tooth. A characteristic feature of the skeleton is the great flexibility of the spine, and the increased area and special form of the articular surfaces of the phalanges and their position in relation to each other, which allows for retraction and extension of the claws (see above). Thoracic vertebrae, 13. Os penis rudimentary. The relative length of the intestine, as a result of feeding exclusively on meat, is small; the relative weight of the heart is small, much smaller than in species of the dog family. Felids live in very diverse habitats, ranging from humid tropical forests _ to sandy deserts. They are usually associated to some degree with forest and shrub vegetation (for cover). They inhabit warm or moderately warm climates without a permanent or with a slight snow cover. Only one species (lynx) is regularly associated with areas of snowy winters, either on the plains or in mountains, and another (snow leopard) with high snowy mountains. Part of the range of some species includes regions with a snowy winter (puma, Amur cat, forest wild cat, tiger, and others). All species are predators, mainly of homoiothermal animals of extremely varying size, from small rodents and birds to giraffes and buffaloes. More rarely, they catch reptiles; some are specialized for catching fish. They hunt by lying in ambush along tracks, near water holes and solonetzes, or by concealing themselves and overtaking the quarry in one or more leaps. They are not capable of prolonged running or chasing. Only cheetah can pursue its quarry for some hundreds of meters and overtake even a gazelle, by attaining a speed maximum for mammals.* They are solitary animals and ’Cheetah are usually considered capable of attaining a speed of up to 140 km per hr. According to recent data the maximum confirmed speed is 56 miles (90.12 km) per hr (Роштей, 1964). 56 55 do not form pairs even at the time of raising the young. Evidently only one species (lion) lives in pairs and family groups and hunts in groups. The den is in the open or located in burrows, rock crevices, and caves. Cats are predominantly nocturnal animals. The range of the family is extremely extensive and covers all zoogeographic regions except Australia (Fig. 18). In the New World cats are distributed from the northern coast of the [North American] mainland to Patagonia and the Strait of Magellan. The family is absent on the American Islands in Bering Sea, except Nunivak (?) [sic]. In the Arctic Archipelago the boundary of the range encompasses Baffin Island, Southampton Island, the small Hudson Bay Islands, Newfoundland, Islands of St. Lawrence Strait, and islands along the west coast of North America (Kodiak, Vancouver, Alexander Archipelago, and others). Representatives of this family are absent in the West Indies, including the Bahamas, Cuba, and Trinidad. On the coast of South America, Chile falls within their range. Cats are absent in Tierra del Fuego and the Falkland Islands. In the Old World the range encompasses almost the whole of Europe in the north, to the tree line. In the west the range includes England; members of the family are absent in Ireland and evidently have never been present, at least not in the historic period. In the Mediterranean Sea the range covers Corsica, Sardinia, Sicily, Crete, and Balearic Islands (in the rest of the islands, including Rhodes, Cyprus, and Malta, cats are absent). Members of the family are absent and were never present in islands of the Barents Sea and small islands and archipelagos of the Atlantic Ocean. In Asia the northern boundary of the range runs along the northern tree line, or a bit to the south, including Kamchatka in northeast Siberia. The southern boundary encompasses the Arabian Peninsula, India, Ceylon [Sri Lanka], and running along the coast of Bay of Bengal descends south to encompass Sumatra and Java, extending between Bali and Lombok, encompassing Kalimantan (Borneo), Palawan and some of the Philippine Islands (evidently only Negros and Cebu in the center). In the Mentawai Archipelago (west of Sumatra), and also on the islands of Billiton, Bangka, and Madura (Java Sea), members of the family are absent. Cats are also absent in the small islands south of Singapore, between Sumatra and Kalimantan (Borneo), and in the South China Sea. Farther, the range bound- ary encompasses Hainan and Taiwan, leaving out the Ryukyu and Japanese islands, but includes Tsushima and Cheju Do (Quelpart), encompassing Sakhalin but leaving out the Kurile Islands, and continues toward Kamchatka. Members of the family are absent on Karaginsk and the Commander Islands, and evidently also on the Shanter Islands. The range covers all the African mainland; Madagascar, and Atlantic and Indian Ocean islands are excluded. Cats, especially the large ones, have [-] а | | НН ` ии НРК TH Ч ий va | | № i ul ill || s/s E> о о ЕЕ ЕЕ ee ЕЕ, Е md, G: О о yo). 5 2) У 90 8 | | И =. an mia ea 1 Se a ET Le ЕЕ, 150 50 20 к? СЗ = т = .. =i LO roe bys mH EIS = ys г = 7 5] ‚аа LE FIG ian ло 7 Lan АЕ 92 р he ou у". `ь. ys 8 © a Oz, = д и ес а >. =e а РА ЕС Fae / й— " 7-37 25 J a РР ых a ЕЯ (| = wt Е ra <% = wa = wee & a у = Fig. 18. Reconstructed range of the cat family, Felidae and small cats, genus Felis L. (scale 1 ees in a о С: in km). V.G. Heptner. 57 vanished at some places and the range of the family has shrunk. On the other hand, domestic cats which become feral under certain circumstances, especially in tropical countries, have settled on some islands outside the natural range of the family. Thus Felis megalotis, described as a separate species from Timor (and Roti), in spite of some persistent objections (Weber, 1928), is considered feral.* The cat family, excluding the Malagassy Fossa (Viverridae), is a highly compact natural group. This family is well separated from related families, above all the closely allied civets (Viverridae).° This is true of the family as a whole, i.e., including fossil forms. Even such extreme forms as saber- toothed cats are typical of the family. Attempts to raise some fossil groups, mainly saber-toothed cats, to the status of families (an extreme point of view — Kretzoi, 1929), have not been recognized (Simpson, 1945; Gromova, 1962). Nevertheless, some recent taxonomists have placed saber-toothed cats in their own family (Machairodontidae) (Thenius and Hofer, 1960). Proposals for segregating cheetah and lynx from Recent Felidae into a separate family (Guepardidae Gray, 1869; and Lyncidae Schulze, 1900) are not well justified and have found little support. At present it appears more natural to view the family broadly and accept five subfamilies: extinct Proailurinae Zittel, 1893; Nimravinae Trouessart, 1885; Machairodontinae Gill, 1872; Hyaenaelurinae Pilgrim, 1932; and con- temporary Felinae Trouessart, 1885. Machairodontinae (Lower Oligocene- Pleistocene; about 10 genera) are considered a very specialized group, standing in rank above the Felinae (Simpson, 1945). The more primitive cats of the subfamily Proailurinae (about five genera) existed from Late Eocene (Early Oligocene) through the Middle Pliocene in the Old World. Some of their early forms combine certain features of cats and civets (‘‘structural ancestors’’), indicating the course of development of the family and its affinities. The original forms (‘‘ancestors’’) of present-day cats are recognizable among members of subfamily Nimravinae (Upper Eocene-Lower Pliocene of Eurasia, North America, and Africa; about nine genera). Fossil forms of the contemporary subfamily Felinae are known from Asia and in the Pleistocene ‘This form is neither mentioned in the most recent catalogs of the family nor in the faunal compilations (Laurie and Hill, 1954, for example). The zoogeographically exceptional case related to the possible occurrence of species in the family in the Australian zoogeographic region has not been discussed. “Its affinity with civets is plainly evident, however. Civets of genera Prionodon and Pardictis (linsangs of southeast Asia) have a short penis, characteristic of cats, which adjoins the scrotum; they have no perineal glands and the claws are retractile and covered with a dermal sheath, another characteristic of cats. The isolated position of family Felidae is emphasized by the fact that cats differ sharply in serological characteristics from other predators. The range of these differences is greater than differences between some carnivores and all pinnipeds (V.I. Borisov). 58 (perhaps Pliocene) of North and South America. Apart from various species of the genus Felis, two fossil Asiatic Lower Pliocene genera (Sivaelurus Pilgr. and Dinofelis Zdansky) and one Pleistocene genus (Sivapanthera Kretzoi) have been described. Certain aspects of the taxonomy of present-day cats are far from simple, and confusion and complications persist even today. This applies to supra- specific groups (the question of the species of cats is now clear). At present, there is hardly a family of present-day mammals in which several conflicting viewpoints relating to their systematics cannot be adduced. Thus the classification of present-day cats into two subfamilies—cheetah (Acinonychinae) and the remaining species (Felinae)—long supported, was replaced by three subfamilies by separating Pantherinae, or “‘big’’ cats, from Felinae (Pocock, 1917; Ognev, 1935; and many others). A later suggestion classified present-day cats into four subfamilies by further separating lynx (Lyncinae) from true cats (Felinae) (Haltenorth, 1957; Weigel, 1961; Mazak, 1965; and others). An extreme view, which however evoked no support, proposed the classification of cats into 15 subfamilies. A fifth subfamily consisting only of the clouded leopard Neofelinae with one genus and species has also been proposed (Kretzoi, 1929). The views of various present-day taxonomists on the classification of cats at the level of subfamily are highly diverse. Most propose two (cheetah, Acinonychinae, and all other cats, Felinae) or three (cheetah, big cats, Pantherinae, and the remainder). Some recognize four subfamilies, establishing a separate subfamily for lynx (Lyncinae). The position of individual species thereby becomes extremely diverse: clouded leopard (F. nebulosa) appears sometimes under Pantherinae and sometimes outside this group; manul is sometimes placed among Felinae and stands close to sand cat (F. margarita) or sometimes in the lynx subfamily (Lyncinae). The latter case exemplifies well a formal approach to taxonomy, disrupting its naturalness. In all craniological and other characteristics the manul is a close relative of the sand cat (see below in systematic section) and represents an extreme degree of specialization among ‘‘small cats”’ in particular and contemporary cats in general. However, the manul lacks the anterior upper premolar (р?) and, based on this single formal characteristic, it falls into the nearest genus, in the same group with lynxes, with which the manul has hardly anything in common. The classification of subfamilies in other cases is even less satisfactory. Thus, a comparison of the diagnostic features of the subfamilies of small cats (Felinae), big cats (Pantherinae), and lynx (Lyncinae) (see for example, Mazak, 1965) reveals that, in the ultimate analysis, the only diagnostic feature of Lyncinae is the tufts on the ears. Quite evidently, this feature is not adequate enough for its separation. 59 The inclusion of big cats, i.e., lion, tiger, leopard, and jaguar, usually also the snow leopard, and clouded leopard (Neofelis)® in a separate subfamily (Pantherinae), which at first appears quite natural, is also unacceptable. There are no sharp differences between this group and Felinae. Insofar as the clouded leopard is concerned, it has no features which are characteristic of Pantherinae proper, i.e., lion, tiger, leopard, and jaguar, while the snow leopard exhibits features of both ‘‘Pantherinae’’ and ‘‘Felinae’’ in an almost equal ratio (Hammer, 1967; see also characteristics of the genus below). Thus there is no distinct demarcation even between typical big cats (“‘Pantherinae’’ — Поп, leopard, jaguar, and tiger) and small cats (“‘Felinae’’), at the subfamily level. The snow leopard is a connecting link between these groups; it is customary to consider its features inter- mediate (see description). Sometimes contemporary cats are not classified into subfamilies; instead all are placed in one subfamily (Felinae) as opposed to the extinct species (Ellerman and Morrison-Scott, 1951 and 1966; Ellerman, Morrison-Scott and Hayman, 1953). This viewpoint reflects more correctly the natural relations of the species of the family but is hardly a prevalent one. It is also my opinion thata clear-cut separation of even genera of contemporary cats is not only difficult but almost impossible (see characteristics of genera below). This is true of even the cheetah; the features of which always call attention to it. The actual differences in this genus are fewer than generally considered (see characteristics of the genus below). Thus, only one subfamily can be identified among contemporary cats.’ The situation in the generic groupings of present-day cats is chaotic. This is due not so much to the complexity of the problem itself, as confusion in the concept of genus in modern taxonomy, primarily due to the aversion of some specialists in systematics for large genera and their attempts to create genera with fairly equal numbers of species.’ Thus a general tendency to ®In spite of its large size, the puma exhibits no affinities with the group of big cats (genus Panthera, **‘Pantherinae’’), a fact long recognized even by advocates holding extremely disparate views. Although this animal is large, it is a true small cat (Felis) in all morphological features. Herein we do not place puma among big cats (whatever be the interpretation of the group) and have no intention of doing so at any time. The number of works devoted to the supraspecific taxonomy of cats is very large. The work of N.A. Severtsov (1858) is the foremost and most interesting study; it exerted great influence on all subsequent works and enjoys importance even today. The works of Pocock (1917) and Haltenorth (1936 and 1937) are also significant. At present, several works have been published which are essentially morphological; even ethological works have been published in recent years (Weber, 1928; Kretzoi, 1929; Tsarapkin, 1932; Ognev, 1935; Pocock, 1939 and 1951; Simpson, 1945; Leyhausen, 1950 and 1956; Haltenorth, 1953; Thenius and Hofer, 1960; Weigel, 1961; Hemmer, 1964; Mazak, 1965; Vereshchagin, 1967; and others). Much of the material on big cats has been reviewed and revised by Hemmer (1966). 8бее Heptner (1965) for “‘larger’’ genera in the structure of systematic groups and their evolutionary significance. 59 60 further divide genera has developed. Cats, in this respect, provide excellent material and there is no other group of mammals in which this tendency has been more manifested, resulting in many major contradictions and distortions. Yet there are some contemporary taxonomists who project a fairly natural picture of supraspecific groupings of cats. All of them propose a very small number of genera. In essence, N.A. Severtsov (1858) was the first to make a broad, theoretically based analysis of the supraspecific groupings in the family. He established a total of 5 genera with 27 subgenera: Tigris (lion and tiger), Panthera (jaguar, leopard, ounce, and puma—four subgenera), Cynailurus (cheetah), Lynchus (lynxes and caracal—two subgenera), and Felis (all remaining cats—17 subgenera). Severtsov’s system, which by and large is well substantiated and thought through, remains unsurpassed even today. . Pocock (1917) considered Severtsov’s classification basic, from one of the modern viewpoints. However, Severtsov’s concepts were altered greatly in principle by Pocock in that most of the subgenera proposed by the former author, primarily in the genus Felis, were raised to the rank of genus. Thus Severtsov’s natural genus Felis, which is “‘large’’ by modern standards was downgraded. Pocock (1917) divided the family into 17 genera ard subgenera. Some contemporary theoreticians have split these genera arbitrarily or follow approximately the same combination. Thus, some accept (while maintaining 3 or 5 subfamilies) 14 (Haltenorth, 1957), 19 (Weigel, 1961), 20 (Mazak, 1965), and even 24 (Ognev, 1935) genera. This system of classification is dubious even on theoretical grounds. According to Mazak (1965), of the 20 genera, 13 (65%) are monotypic, 1 contains 2 species, 3 contain 3 species each, 1 contains 4 species, and another 5 species. With 24 genera (Ognev, 1935), the number of monotypic genera rises even higher. Thus generic characteristics, features, and the very concept per se become species-based. The natural concept of a monotypic genus in such a case is largely lost. In point of fact it appears at times that between some species within the same genus differences are incomparably larger and more significant (Mazak, 1965) than differences between genera (Felis—Prionailurus; Lynx—Caracal; and so on). Such a situation is arbitrary and difficult to comprehend. Various authors not only evaluate differently the rank of individual forms (big cats—lion, tiger, leopard, and jaguar—one genus or four monotypic genera, and so on), but they also occupy very different positions in the series (evolutionary, specialization) of forms in the family. Thus, the manul is placed as a distinctive genus sometimes in the immediate proximity of genus Felis (Mazak, 1965), sometimes there are 14 genera and 7 subgenera intervening between them and it is placed in a separate ‘‘subfamily,’’ i.e., lynx (Weigel, 1961), and so on. All this underscores the 61 artificiality of excessive subdivision of contemporary members of the family into genera. An opposite, extreme viewpoint recognizes only two genera—Acinonyx (cheetah) on the one side and Felis, which includes all remaining contemporary cats, on the other. Usually the latter genus is extensively subdivided into subgenera. This system, which is quite reasonable (see later), has several supporters (Allen, 1938; Novikov, 1956; Hall and Kelson, 1959; and others). Yet in spite of all the diversity of size and shape of individual species, large craniological differences, and relatively large number of species (35), contemporary cats represent an extremely homogeneous group morphologi- cally, systematically, and phylogenetically. In practice, the position is that almost any two species which are extremely dissimilar can be linked through transitional forms into a fairly continuous morphological series. Breaking this chain or mosaic pattern into innumerable independent genera is almost invariably artificial. Even the characteristics of the cheetah (see below) are less sharp and more typical than usually admitted. This applies to behavioral characteristics also (Leyhausen, 1950 and 1956; Hemmer, 1964 and 1966; and others). It is interesting that in [almost] all species, as far as is known, the number of chromosomes is identical (2n=38) and the differences in karyotype insignificant; only two (F. pardalis and F. wiedi) have 36 chromosomes* (V.N. Orlov). Under artificial conditions (zoological gardens) all species of large cats have produced hybrids (lion, tiger, leopard, and jaguar) and even trihybrids; backcrosses are also successful. So far only in the case of the snow leopard are hybrids not known; however, it does not breed readily in captivity; and tiger with leopard (information relating to such hybrids in nature is not reliable). But then, hybrids have been produced by crossing leopard with puma, which is not a big cat (genus Panthera), 1.е., a case of intergenus hybrid (Hemmer, 1966). The diversity of interpretation of the above-noted characteristics of the family notwithstanding, natural groups are known which, according to some other possible grouping of the species, could be assigned the rank of genus among present-day cats. Three or four such groups are recognized. According to one point of view (Simpson, 1945), the family consists of three genera— big cats, Panthera (lion, leopard, tiger, jaguar, snow leopard, and clouded leopard), cheetah (Acinonyx) and small cats (Felis) including all the remain- ing species of the family. Here the subfamilies of Pocock (1917) are treated as genera. According to an identical viewpoint (Ellerman and Morrison- Scott, 1951 and 1966) the same genera with similar species composition *As of time of publication of Russian edition—Sci. Ed. 60 62 are recognized; however, the clouded leopard (F. nebulosa) is allocated to an independent genus (Neofelis), occupying a position between the genera Panthera and Felis; in all four genera are recognized (Palearctic and India). It is becoming increasingly clear (see the new works cited above, especially that of Hemmer, 1964 and 1966) that Neofelis has no affinities with big cats (Panthera). Neither does it assume an intermediate position between big and small cats (Felis). This is, apparently, a genuine small cat which is evidently no more peculiar than some other more or less ‘‘peripheral”’ species of this group. It is placed here as a member of genus Felis (as a subgenus). On the other hand the snow leopard is not wholly typical of the genus of big cats (Panthera) and combines fairly equally the characteristics of both big (Panthera) and small (Felis) cats.? However, it differs from both, possessing features typically its own, and, in spite of also exhibiting features of an intermediate nature, is separated as an independent genus.!° | Thus, as in the works of other researchers cited above, four genera are recognized here also, but somewhat differently and with differences in species composition: big cats (Panthera Oken, 1816), ounce or snow leopard (Uncia Gray, 1854), small cats (Felis Linnaeus, 1758), and cheetah ог pardus (Acinonyx Brookes, 1828)."! The genus cheetah (Acinonyx) is unique and well separated, albeit less well so than commonly admitted. This genus is close to small cats (Felis) and is undoubtedly derived from them. The cheetah is a specialized cat, the characteristics of which lie in adaptation to fast running, 1.е., in a direction altogether unusual for cats, and to catching relatively large prey. It exhibits no affinities with big cats (Panthera). The demarcation between genera of big cats (Panthera) and small cats (Felis) are far less definite. Significant features of an intermediate character, as pointed out above, are seen in the snow leopard (genus Uncia), which connect them quite clearly. There are also forms of true small cats exhibiting affinities toward the snow leopard (clouded leopard, puma). Evidently these are features of convergence resulting from the large size of the animal as °This situation may provide a base for combining genera of big (Panthera) and small (Felis) cats in а common genus. This approach is possible; however, it applies only to those subgeneric groups which could be separated within the limits of genera Felis and Panthera, in relation to the taxonomic weight of their features. Details of all genera are given later under their description. 'IWith reference to the degree of taxonomic fragmentation of contemporary cats and in the approach to supraspecific groupings in the family, i.e., few genera and extensive use of the concept of subgenus, the views of some contemporary scientists (Ellerman and Morrison-Scott, Simpson, and Heptner) come very close to those of М.А. Severtsov (1858). 61 63 well as its prey. Thus Panthera, Uncia, and Felis form a close group or a chain of successive genera. Relative to the level of organization (“‘primitiveness’’ or “‘progressive- ness’’) of these genera and the degree of their morphological specialization, it is quite difficult to resolve the question of monotypic species in the family and the closeness of their relationships are quite troublesome. One is hampered additionally because paleontology provides very little data due to the fact that various types of present-day cats appeared quite simultaneously. Evidently even the directions of various adaptations among present-day Felinae also proceeded more or less simultaneously. According to the traditional and more prevalent point of view, all small cats of the genus Felis form a least specialized (‘‘primitive’’) group, and big cats of the genera Panthera and Uncia, as well as cheetah, a more specialized (‘‘progressive’’) group. The latter is usually considered specialized in a special sense and in a direction different and independent of big cats. In giving a very general evaluation of the skull structure of small cats of the genus Felis (apart from those such as puma and clouded leopard), attention is drawn to features such as the very large volume and bulging form of the cranium, the very short facial region, and the absence or slight development of various types of crests and tuberosities serving for the attachment of muscles, mainly the group of masticatory muscles. These features are usually considered juvenile (infantile) and often justifiably Fig. 19. Skull of tiger (Panthera tigris Г.) and manul (Felis manu! Pall.) reduced to the same scale, two extreme types of contemporary cats. Collection of the Zoological Museum, Moscow University. Sketch by N.N. Kondakov. 61 64 regarded as an index of the slight specialization of the group, that is to say, more neutral than those of more primitive structure. The opposite type of features characterizing skulls of big cats are considered specialized and hence progressive. This traditional approach is hardly applicable to contemporary cats. The skull of big cats with its highly developed facial region, powerful crests, and relatively moderate, elongated braincase, has an appearance and structure of the skull far more similar to that of predators of other families than does the skull of small cats. The skull of big cats exhibits a greater degree of general predatory features than specific feline characters and structure (Fig. 19). Members of the small cat genus Felis $. 1. exhibit features that are more sharply specific for contemporary cats. The differential characteristics of modern Felidae are more fully manifested in the skull of small cats. Thus progressive specialization within present-day cats proceeded in the direction of the development of a superior feline type, as seen in the manul but not in tiger. In spite of all the specialization of big cats, typical feline characteristics in them are fewer than in small ones and less distinctly manifested. Small cats are truer representatives of cats in the total meaning of the term. In spite of scanty paleontological material, one can conclude that present-day big cats (genus Panthera) exhibit a larger number of features of similarity with fossil forms, even with totally extinct groups, than present- day small cats. Fossil forms characteristic of the Felis type (silvestris, for example, or /ynx), not considering the most recent direct antecedents to contemporary forms, are either unknown or few in number. Evidently the evolution of a fully developed type of contemporary small cat (genus Felis) represents a new stage in the evolution of the family and subfamily. The relative primitive nature of big cats has in fact been noted by paleontologists though, it is true without arguments (Thenius and Hofer, 1960), although usually they support a contrary view (Simpson, 1945). The placement of modern cats with a ‘‘juvenile”’ type of skull with progressive members of the family is unusual, but the situation is not unique in predatory mammals (Ferae). The family of common seals (Phocidae), invariably recognized as the most specialized and progressive in the order (suborder), differs from eared seals (Otariidae) in skull structure. For example, the genus Eumetopias (sea lion) exhibits particularly well- manifested overall predatory features in the skull. The course of development of the family evidently lay in the course of cephalization. Increase in overall size in a phylogenetic series is usually considered a sign of specialization and advancement. It is, however, hardly possible to assign an absolute and decisive importance to increase in size; size is extremely relative in these same pinnipeds. The number of species in the 62 65 genera also underscores the progressiveness of the small cat group (Heptner, 1965): about 11.5% of the species in the family fall in the genus of big cats (Panthera), 3.5% each in the genera of cheetah (Acinonyx) and snow leopard (Uncia), and 81.5% in the genus of small cats. Thus there is adequate justification to begin the phylogenetic series of contemporary cats with big cats (Panthera), followed by snow leopard (Uncia), and then small cats (Felis). The cheetah (Acinonyx) represents a specialized lateral branch of the line of small cats. These groups represent three main directions of specialization with respect to size of prey and method of hunting—catching large animals (big cats and snow leopard), catching small vertebrates (sometimes larger; small cats), and catching large animals by pursuit (cheetah). The family Felidae contains fewer species than civets (Viverridae, about 75 species) and weasels (Mustelidae, about 70 species); nevertheless with 35 species the family comprises 14.5% of the total number of species in the order and about 1.0% of all mammalian species. Felidae outnumbers the dog family species-wise (Canidae, 29 species). In the genus of big cats (Panthera) there are four species; ounce or snow leopard (Uncia), one; small cats (Felis), 28; and cheetah (Acinonyx), one. Of the four genera, one (cheetah) is Afro-Asian; one (Panthera) mainly Old World with one representative (jaguar) in South, Central, and North America (extreme southern part of North America*); one (Uncia) wholly Asian; and one (Felis) widely distributed throughout the range of the family in the Old as well as New World. The maximum number of species are associated with the Old World, particularly Asia; the smallest number inhabit North America; a fairly significant number are found in South America. The economic importance of the family as a whole is minimal and depends on the level of economy in a given region. In untouched biocenoses and regions little settled by man, species of the family have little or no practical importance as fur animals or in the trade of live animals for zoological gardens. In more densely populated regions big cats often kill cattle or damage game prospects by killing wild ungulates. Some individual animals (tiger, lion, sometimes leopard and jaguar) pose a danger for man under certain conditions in a given region. Some large species (lion, tiger, leopard, puma, and jaguar) are valued as game and sport animals. At the same time they exert a selective influence on wild ungulated populations, which is a positive factor. On the other hand, in cultivated areas even small cats may damage the hunting economy by destroying game birds (pheasants), hares, or young ungulates. Many species feed almost exclusively on small rodents but are seldom effective in *Closing parenthesis out of place in Russian original —Sci. Ed. 63 66 controlling rodent populations. Some species are protected in certain areas or sanctuaries. One (F. silvestris) also survives as a truly domesticated animal (domestic cat, ‘К. catus L.’’)!? and two species have been tamed and trained for hunting (cheetah for hunting gazelle, and caracal for hunting birds); but cannot be considered ‘‘domesticated.’’ In all, 4 genera and 11 species, i.e., about 31 to 32% of species of family Felidae, are found in the Soviet Union. They comprise about 10% of the species of the Russian fauna. The range of Felidae covers almost all of the northern USSR to the tree line. The majority of species are associated with the southern regions of the country, however; only one (lynx) also occupies extensive regions in the north. There are forest, montane, steppe, and desert forms. They are of relatively moderate significance as fur animals or as exhibits for zoological parks. Due to persecution the population of some species has fallen sharply and these are now on the verge of extinction. One species (tiger) has suffered a particularly dramatic reduction in population and range and is placed under conservation; two other species (cheetah and caracal) are under protection. (V.H.) Key for Identification of Species of the Cat Family, Felidae'> Identification by External Characters — ( 2). Sharp, vivid, dark (black) band extends from inner corner of eye along nose toward upper lip on a light field. General color very light, yellowish with small, black, dense, circular spots not forming rings. Claws project significantly from fur covering paw ...... а. cheetah [gepard]* Acinonyx jubatus (р. 702). ( 1). Well-developed black band extending from inner corner of eye to upper lip absent. General color different. Claws do not project from fur covering paw. 3 ( 4). Vivid vertical (transverse) black stripes extend through bright reddish background color on back and flanks. Very similar pattern of stripes occurs on head. Size very large (body length of adults exceeds 60cm) о. tiger [tigr] Panthera tigris (p. 95). 4 ( 3). Color different. Size smaller. 5 ( 8). Groups of black spots arranged in form of rings dispersed throughout light background together with individual dense spots iw) Currently labeled Felis silvestris L. forma catus. See the section **Description’’ under steppe cat (К. 5. caudata) for the origin of domestic cats. '3Since the lion is no longer found in the Soviet Union, it is not included in the key. *Transliteration of Russian common names in brackets. 61). 7 ( 6). 8 ( 5). 9 (10). 10 ( 9). i 2). 1D В 13 (16). 14 (15). 64 15 (14). 16 (13). 67 of moderate size. Size fairly large (body length of adults exceeds 90 cm, often over 100 cm, and even 120 cm). Main color of trunk light yellow or reddish yellow. Tail length less than length of head and body. Spots sharply outlined and vivid black . . . . leopard [bars, ‘Чеорага’”] Panthera pardus (р. 203). Main color not yellow but dirty white or very light gray. Tail length roughly equal to length of head and body. Spots, at least in winter coat nobvividsand not Sharply,promled S22)... 2 2 cs es snow leopard, ounce [snezhnyi bars, irbis] Uncia uncia (p. 276). Color different. If spots present not arranged in form of large rings. Tail length, without terminal hair, less than that of hind foot or equal to it; terminal one-third or one-fourth of tail black ....... Bee Niet a capers We i es So aia lynx [rys’] Felis lynx (p. 524). Tail length, without terminal hair, more than length of hind foot. Tail end not black or black for less than one-third or one-fourth of tail length. Two distinct light (white) bands, usually bordered by black, extend from inner corners of eyes along forehead. Back of ears with well- defined white field (spot). Ocherous or reddish spots or transverse band visible on throat and lower surface of neck ........... . .. leopard cat, Amur cat [amurskii kot] Е. euptilura (р. 328). White bands extending from inner corners of eyes along forehead absent; rear of ear without white spot; reddish or ocherous spots and transverse bands on throat and lower surface of neck absent. Spots distinctly visible. General tone of back and flanks brownish-gray. Along back, double line of merged spots visible and similar vertical bands present on sides in addition to individual spots. Throughout its length tail uniformly covered with long hair (not thinned toward the end, terminal hair long) and appears trimmed. Tail with several (3 to 10, usually 5 to 7) black transverse bands (Fig. 20)*..... forest wild cat [Evropeiskaya lesnaya koshkal], Felis silvestris, group silvestris (p. 402). General tone light dirty gray or pale gray. Back and flanks covered with innumerable distinct spots. Double black band along back and transverse stripes on flanks not well defined, or very faint. Raili\tapers toward ends (Fig, 20)... steppe wild cat [stepnoi kot], Felis silvestris, group libyca (р. 441). Body monochromatic, without distinct spots, or with several dark, narrow, transverse bands running across back. Dark marks (spots *Erroneously given as 19 in Russian original—Sci. Ed. Fig. 20. Form and pattern of tail of European (and Caucasian) forest wildcat, Felis (Felis) silvestris, group silvestris and steppe wildcat, Е. (F.) silvestris, group libyca. Collection of the Zoological Museum, Moscow University. Sketch by N.N. Kondakov. or bands) may occur on legs; if some individual spots occur on body, they are blurred, indistinct, and small. 17 (18). Back of ears black; tuft of long (usually over 20 mm) hair at tip И Е в. caracal [karakal], Felis caracal (р. 498). 18 (17). Back of ears not black; tuft on ears absent, or hair on tip of ears shorter (less than 20 mm). 19 (20). Lower surface of paw with long, dense, black or blackish-brown hair covering sole and digital pads ....................0-. и sand cat [barkhanaya koshka] Felis margarita (р. 636). 20 (19).*Lower surface of paw devoid of long, dense, black hair and sole and digital pads hairless. 21 (22).*Tail relatively short and does not, or barely reaches calcaneal joint. Long hair on cheeks (whiskers) absent. Hair at tip of ears sometimes slightly elongated. Ears large, with pointed tips, and project beyond fur imwinter и = jungle cat [kamyshovyi kot] Felis chaus (р. 356). 22 (21).*Tail relatively long, end extending significantly beyond calcaneal *Misnumbered in Russian original— General Editor. 69 joint. Long hair (whiskers) present on cheeks. Hair at tip of ear not elongated. Back of ear with blurred, narrow, transverse bands. Ears short, rounded above, and project very little from dense rich coat in winter. . , . manul cat [manul], Felis manul (р. 655) (V.H.) Identification by Skull Characters'4 . Size large. Length of upper carnassial tooth more than 21 mm. . Length of upper carnassial tooth 29 mm or more. Condylobasal length 250 mm ormore...... tiger, Panthera tigris (p. 103). . Length of upper carnassial tooth less than 29 mm. Condylobasal length not more than 240 mm. . Maximum width of interpterygoid vacuity more than least distance between tympanic bullae. Inner cusp of upper carnassial tooth not developed or barely visible and devoid of peak (tubercle) (Fig. 21). Distinct small cusp present in front of main cusp of third premolar (before Camassial)) (Ее 22)... Е orleans cheetah, Acinonyx* jubatus (p. 702). Fig. 21. Upper carnassial tooth of cheetah, A cinonyx jubatus (left) and snow leopard, Uncia uncia. Nos. S 47284 and S 49240, collection of Zoological Museum, Moscow University. Sketch by N.N. Kondakov. 140 п examination of material some of the ‘‘traditional’’ features used for the identification of this genus were found to be inconsistent; namely, form of infraorbital foramen was sometimes, and in some species often, double; most features of relative disposition of posterior end of nasals and frontal processes of upper jaw; condylobasal length; and other major features, viz., position of anterodorsal end of jugal in relation to lacrimal duct. In the key given here an attempt has been made to include some new features together with some fairly common ones. 1SApplies to permanent and not deciduous tooth. Reliable identification of juveniles, especially from deciduous dentition, is possible only in museums with significant comparative material. *Misnumbered/misspelled in Russian original — Sci. Ed. 66 70 Fig. 22. Third and fourth (camassial) premolars of snow leopard, Uncia uncia (top) and cheetah, Acinonyx jubatus. Nos. S 47284 and S 49240, collection of Zoological Museum, Moscow University. Sketch by N.N. Kondakov. 54): Maximum width of interpterygoid vacuity less than least distance between tympanic bullae. Inner cusp of upper carnassial tooth well developed and with prominent peak (tubercle) (Fig. 21). Distinct cusp in front of main cusp of third premolar (before carnassial) absent (Fig. 22). 6 ( 7). End of paroccipital process extended, overhanging rear of tympanic ТСО. a @ 1): bulla in the form of very faint pointed hook (Fig. 23*). Antero- lateral (ectotympanic) chamber of tympanic bulla well developed (Fig. 23). Anterior wall of swollen portion of bulla almost at level of posterior wall of glenoid fossa (postglenoid process; Fig. 23) т snow leopard, Uncia uncia (р. 284). Paroccipital process short, thick, and blunt; adjoins posterior wall of tympanic bulla completely; no long, thin, overhanging end in form of a hook (Fig. 23). Almost all of bulla consists of postero- medial (entotympanic) chamber; anterolateral (ectotympanic) portion very small (Fig. 23). Anterior wall of swollen portion of bulla separated from posterior wall of glenoid fossa (postglenoid process) by a distance roughly equal to width of buila (Fig. 23) Wala aw eats oly SEN eon cues leopard, Panthera pardus (p. 208). Size small. Length of upper carnassial tooth less than 21 mm. 9 (10). Length of upper carnassial tooth 17 mm or more.!© Second upper *Misnumbered in Russian original — Sci. Ed. 16 п rare cases, when length of upper carnassial tooth is about 17 mm (+ 0.1 to 0.2 mm), the characters should be compared with those given in parentheses in 15 (16). 71 ме ВИ о ВАМ ААА С N х Е | Ri ’ Fig. 23. Tympanic bulla and paroccipital process in snow leopard, Uncia uncia (left) and leopard, Panthera pardus. Upper row—ventral view; lower row— lateral. Nos. S 35132 (snow leopard) and S 51945, collection of the Zoological Museum, Moscow University. Sketch by N.N. Kondakov. premolar (second tooth before carnassial) absent (three teeth occur in upper jaw behind canine).!’...... lynx, Felis lynx (p. 532). 10 ¢ 9). Length of upper carnassial tooth less than 17 mm. Second premolar in upper jaw present or absent (three or four teeth occur beyond canine). 11 (14). Ectotympanic chamber of tympanic bulla very large; bullae greatly swollen and very large; its anterior point at level of posterior wall 1ТА$ a rare exception, very small second premolars may be present but usually only on one side. 66, Fig. 24. Tympanic bullae of manul, Felis manul (left) and sand cat, F. margarita (center) with highly developed anterolateral (ectotympanic) chamber. Right, for comparison, bulla of steppe wildcat F. silvestris, group libyca, with a very small anterior chamber. Nos. S 40113, S 64765 and S 42325, collection of the Zoological Museum, Moscow University. Sketch by N.N. Kondakov. of glenoid fossa (postglenoid process) or even slightly shifted forward (Fig. 24). 67 12 (13). Inner cusp of upper carnassial tooth well developed and with distinct small peak (tubercle; Fig. 25). Maximum diameter of external auditory meatus roughly equal to length of upper camassial tooth. Second premolar present in upper jaw (four teeth behindicanine) 33 о sand cat, Felis margarita (p. 642). 13 (12). Inner cusp of upper carnassial tooth not developed and peaks (tubercles) not evident (Fig. 25). Maximum diameter of external auditory meatus less than length of upper carnassial tooth. Second premolar on upper jaw (three teeth behind canine) ........ DE SE PEERS cicero ce as manul cat, Felis manul (p. 671). 14 (11). Anterolateral (ectotympanic) chamber of tympanic bulla small and almost all of bulla formed by posteromedial (entotympanic) chamber. Anterior point of tympanic bulla does not reach level of posterior wall of glenoid fossa (postglenoid process). 15 (16). Distance between tips of supraorbital processes only twice the interorbital width. Distance between supraorbital process of frontal 67 Fig. 25. Upper carnassial tooth of manul, Felis manul (left) and sand cat, F. margarita. Nos. $ 40113 and $ 67765, collection of the Zoological Museum, Moscow University. Sketch by N.N. Kondakov. 18ботентез this tooth, especially in old animals, is lost from one or both sides but fairly distinct traces of its alveolus invariably remain. 67 16 (15). Fig. 26. Shape of nasal process of premaxilla and its relation to nasal process of frontal in caracal, Felis caracal (left) and Caucasian wildcat, F. 5. caucasica. Nos. S 40195 and S 17640, collection of the Zoological Museum, Moscow _ University. Sketch by М.М. Kondakov. 18 process of premaxilla narrow and long, and tapers gradually upward; tip elongated and sharp; extends closer to forward projection of frontal; distance between them usually not more than length of third upper premolar (Fig. 26*). Second upper premolar usually absent. (A distinct bony crest runs to orbit along outer side of pterygoid bone. At its posteriormost end it forms a broad plate in the form of a triangle or fork. Length of tympanic bulla measured from canal [foramen] of carotid artery to the point of its union with the lateral occipital process is equal to distance between ends of upper canines or slightly more.)...... о cs Ma НО: liad che caracal, Felis caracal (p. 503). Distance between ends of supraorbital processes more than two times interorbital width. Orbit closed from rear or distance between supraorbital process of frontal and postorbital process of jugal less than length of latter. Nasal process of premaxillae relatively short, broad along much of its length, and constricted upward in form of a relatively blunt wedge. Distance between its tip and tip of forward projection of frontal more than length of third upper premolar (Fig. 26). Second upper premolar usually well developed. Posterior margin of bony palate outside interpterygoid vacuity notched, steep, and deep. *Misnumbered in Russian original — Sci. Ed. 1°See footnote No. 16 above. 74 Fig. 27. Shape of nasals in jungle cat, Felis chaus (left) and steppe cat, Felis silvestris, group libyca. Nos. S 3603 and S 42325, collection of the Zoological Museum, Moscow University. Sketch by N.N. Kondakov. 17 (18). Length of nasal, measured from posterior end to anterior projection, more than distance from posterior margin of anterior palatine foramen to posterior margin of bony palate at a corresponding point in interpterygoid vacuity, or exceptionally, equal to it. Posterior ends of nasals usually have an extended, pointed shape A UT DES SS jungle cat, Felis chaus (р. 362*). Fig. 28. Position of posterior margin of bony palate in interpterygoid vacuity in Amur cat, Felis euptilura (left) and European (Caucasian) wildcat, Felis silvestris. Nos. S 14261 and S 75098, collection of Zoological Museum, Moscow University. Sketch by N.N. Kondakov. *Misnumbered in Russian original— Sci. Ed. 75 18 (17). Length of nasal, measured from posterior end to anterior projection, less than distance from posterior margin of anterior palatine foramen to posterior margin of bony palate at a corresponding point in interpterygoid vacuity, or exceptionally equal to it. Posterior ends of nasals have a tapered, only slightly extended shape (Fig. 27). 19 (20). Bony palate long; its posterior margin in region of interpterygoid vacuity lies at level of posterior surface of molars at a distance roughly equal to length of third premolar or at a distance roughly equal to one-half or more of width of interpterygoid vacuity (Fig. 28). Interpterygoid vacuity of uniform width and narrow;”* its inner width in anterior part not more than 10 mm and width along outer walls at level of anterior margin less thaniS mm........... а Amur cat, Felis euptilura (р. 335). 20 (21). Bony palate short; in region of pterygoid [sic] vacuity its posterior margin lies at level of posterior surface of molars, or only slightly beyond this line, at a distance much less than length of third premolar and less than one-half of width of interpterygoid vacuity (Fig. 28). Interpterygoid vacuity somewhat broader anteriorly than posteriorly; broad,* its inner width in anterior part exceeds 10 mm and width along outer walls, at level of anterior margin, exceeds 15 mm. 21 (22). Maximum diameter of external auditory meatus more than length of third upper premolar (lies before carnassial tooth) (Fig. 29)... ........Steppe wildcat, Felis silvestris, group libyca (р. 450). 22 (21). Maximum diameter of external auditory meatus less than length of third upper premolar or equal to it (Fig. 29)............ forest wildcat, Felis silvestris, group silvestris (p. 407). (V.H.) Fig. 29. Tympanic bulla and external auditory meatus of steppe wildcat, Felis silvestris, group libyca (left) and European (Caucasian) forest wildcat, F. s., group silvestris. Nos. S 42325 and S 17640, collection of the Zoological Museum, Moscow University. Sketch by N.N. Kondakov. * Appears inconsistent with Fig. 28, but not Figs. 158 and 183 — Sci. Ed. 76 1816. 1816. 1816. 1829. 1843. 1868. Genus of Large Cats Genus Panthera Oken, 1816 Рапйега.?° Oken. Lehrb. Naturgesch., 3, 2, 1052. Felis pardus Linnaeus. Tigris, Oken. Ibid., 3, 2, 1016. Felis tigris Linnaeus. Leo. Oken. Ibid., 3, 2, 1070. Felis leo Linnaeus. Leo. Brehm. 1515 (ОКеп), р. 637. Felis leo Linnaeus. Tigris. Gray. List Mamm. Brit. Mus., p. 40. Felis tigris Linnaeus. Pardus. Fitzinger. Sitz.-Ber. K. Ak. Wiss. Wien, 58, 1, 459. Felis pardus Linnaeus. (V.H.) Large to very large animals; largest in the family. 70 Fig. 30. Body contour of big cat of genus Panthera and small cat of genus Felis. Elevation of sacral region in small cats is characteristic. Depicted — а lioness and a domestic cat. Sketch by N.N. Kondakov. 20This name does not fully satisfy the rules of nomenclature but is extensively used. It should be retained as a nomen conservandum. 71 77 General appearance wholly typical for all cats. Legs in most not long, but thick; trunk extended, sometimes highly so. Sacral region not high and back not raised toward rear; either straight or height at shoulder (withers) more than at sacrum or back slopes gently toward rump (Fig. 30). Tail long, not less than one-half of body length, or slightly more. Head moderate in size or large, somewhat extended with a relatively elongated facial region, and fairly straight or faintly arcuate dorsal profile. Ears relatively small, short with a blunt rounded tip, and wide-set. In male lions and animals in winter coat ears project slightly out of coat. Long hair (tufts) on top ‚ of ear absent. Eyes relatively small and pupil round. Pelage in most species uniform throughout body, and tail uniformly furred throughout its length. Sometimes hair on cheeks elongated (whiskers), in one species (lion) well-developed mane covers anterior portion of body and tail with long tuft at end. Legs strong with broad, powerful paws, especially forepaws; claws large, very sharp, sharply bent, and fully retractile. At end of all digits, dermal thickening occurs above and below claws; this thickening completely conceals retracted claw. Interdigital membranes large and reach end of digital pads (Fig. 278). In most species hair short and not dense, becoming dense and relatively long only in extreme northern forms of some species in winter. Color monochromatic, with black transverse stripes, or with black spots dispersed against light-colored background as single spots or groups forming rosettes. In most species large white spot occurs on back of ear. Teats, two or three pairs. Skull large and heavy, with massive bones, thick and powerful zygomatic arches, and powerfully developed tuberosities and crests. Sagittal crest particularly long, very high and projects notably backward; lambdoidal crest well developed. Dorsal profile fairly straight or only faintly and uniformly convex; interorbital region raised relatively weakly; line of profile slopes gently back from it. On the whole skull appears flattened. Basicranial axis (floor of cranium) almost horizontal and does not form a distinct angle with basifacial region. Depression (inflexion) at point of transition of nasal into interorbital region on upper part of skull absent. Brain case of relatively moderate size, very small in some, and elongated as though greatly pressed by masticatory muscle attached to sagittal crest. Postorbital constriction marked; sometimes its width is equal or even less than width of interorbital constriction. Facial region of skull very powerful and broad (in region of zygomatic arches and in region of canines) and also elongated. Measuring along a transverse line passing through the supraorbital processes, facial region larger, or not less than, braincase as a whole, or without portion of sagittal crest extending beyond occiput. There is fairly large separation, roughly equal to width of tympanic bulla, 70 78 between postglenoid process and bulla. Tympanic bullae relatively small, not bulging, flattened from ventral side, and thick-walled (small in leopard, but stands out); inner chamber (entotympanic) significantly larger than outer (ectotympanic or tympanum proper), and septum between them lies close to outer ear passage. In hyoid apparatus (Fig. 31) third upper element (ephihyal) not ossified and forms a hinge ligament which permits freer movement of the larynx (see below). Deep depressions absent or barely perceptible on basioccipital portion of occipital bone close to tympanic bulla in front of jugular foramen. Dentition complete; second premolar (first in row) usually present on upper jaw. Teeth very powerful; canines relatively short but strong with a broad base. Sexual dimorphism in most species manifested in large size of males and in part more intense development of crests, etc. on skull; age-related color dimorphism not evident. In one species (lion) male differs sharply in appearance in growth of mane on anterior part of trunk and elongated hair at some places on chest and abdomen; adults monochromatic but color of newborn and young spotted. Seasonal dimorphism in color absent; winter coat denser and with long hair in northern or hill subspecies of some species. These predators feed mainly on big mammals (ungulates). The prey is usually bigger than the predator, sometimes by several times. Hunting is Fig. 31. Hyoid apparatus (sketch) in genus of big cats, Panthera Oken (left), and genus of small cats, Felis L. and cheetah, Acinonyx. Sketch by N.N. Kondakov from H. Hemmer. 71 79 done by concealment and ambush on pathways, at water holes, and so on. They are nocturnal animals, and with one exception solitary. The male does not participate in the feeding of the young. One species (lion) lives and hunts in small groups, representing a family unit, as in the case of wolves (see Volume II, Part 1). They are mostly inhabitants of plains and montane forests, sometimes open forestless mountains, and reed thickets; one species lives in open steppes (savannas) and semideserts. As a result of the specialized nature of the hyoid apparatus (see above), the larynx and vocal chords are highly mobile and the voice a loud coarse 72 roar (‘‘roaring cats’’; big cats are incapable of ‘“‘meowing’’). They do not express contentment and well-being by “‘purring’’, or “‘purr’’ only on exhaling. They mainly eat while resting on the abdomen, keeping the forearm and ulnar region in contact with the ground, and tear the prey to pieces by jerking the head upward (Fig. 32). The range of the genus (reconstructed) (Fig. 33) includes Africa, extreme southeastern Europe, Asia (except the northern part), and South, Central and southernmost North America. In South America the range covers all of the northern and partly the central part of the mainland; southward it includes Peru, Bolivia, Paraguay, and southern Brazil roughly up to Sao Paulo, the province of San Luis, and even La Pampa in Argentina (slightly north of 40° NL). Northward the range encompasses all of Central America, Fig. 32. Usual posture of a big cat of genus Panthera while eating (forearm region horizontal to substratum) and small cat of genus Felis (ulna raised high). Sketch by N.N. Kondakov. Be ь а a al | a | Е oe | и 2 | м ‹ ‘Ny i Sa | Fig. 33. Reconstructed range of big cats, Panthera (scale in as И | | и 1 | mut ey 5. — 30. 0 30 60 90 120 tructed range of big cats, Pantl (scale in km). V.G. Heptner. Е 73 > &y roughly up to the latitude of Veracruz in southern Mexico; and farther, bypassing on the east and west the Mexican highlands, extends along the lowlands of the Pacific Ocean and the Caribbean Sea, and extends in the form of two long extensions into southern Texas on one side and California, Arizona, and New Mexico on the other. The California Peninsula [1.е., Baja California] proper and all islands of the western hemisphere fal! outside the range. The range encompasses all of Africa and the southern parts of the Balkan . Peninsula in Europe. In Asia the southern boundary encompasses the Arabian Peninsula, runs along the southern edge of the mainland, and covers Sri Lanka, Malacca, Sumatra, Java, Bali, and Kangean Island (north of Bali). In the east the boundary similarly runs along the coast of the mainland but species of the genus are not found in Hainan,?! Taiwan, or the islands of Japan and Shanter Islands. In Sakhalin only stray animals have been reported (tiger). In northern Asia the range includes the Amur, Trans-Baikal, Altai, and Kashgar, eastern and southern Kazakhstan, and Middle Asia, Iran, the Caucasus, and Asia Minor (for details see distribution of tiger and leopard). Over the historic period the range, at places, has shrunk due to extinction (Europe, Middle Asia, Siberia, parts of Africa and southern Asia, and others) and continues to shrink even more rapidly elsewhere. Species of this genus are among the best studied in the family, especially their morphology and ethology.?? Differences of opinion among syste- matists relate only to the relative progressiveness of individual species and their division into subgenera (see below). As already mentioned above, there is justification to regard big cats as less specialized among genera of the family. In any case, compared with other groups of the family (different groups of species in genus Felis) in which feline type specialization has reached utmost development and expression, members of this genus represent a less specialized, more generalized ‘‘ргедаюгу’” type in skull structure. It is possible that some features of species of genus Panthera are secondary (allometric). Big cats appeared in the Early Pliocene of Eurasia. In the Early Pleistocene of Eurasia they were represented by special species (P. arvernensis and P. cristata), sometimes considered the ancestral forms of 21У. Mazak (1965, р. 33) has indicated (arrow) on the distribution map of tiger he presents, a sighting on Hainan. This is clearly erroneous. Mazak evidently had in mind old information about the intrusion of tiger from the mainland onto Amoy Island (5 miles from the mainland) cited by Allen (1938). There is no positive reference whatsoever in literature (including also Mazak, 1965) to Hainan. *2Literature devoted to this group of cats is extensive. The basic works are those of Severtsov, 1858; Pocock, 1917; and others; Haltenorth, 1936 and 1937; and others; Weigel, 1961; and others; and the bibliographic compilation of Hemmer, 1966. “as 74 82 lion and tiger. The cave lion or “‘tiger-lion’’ (Р. spelea), extensively distributed throughout the whole of the Pleistocene period, is an independent species with which present-day lion and tiger have no direct genetic association. In the New World the genus is known only in the Pleistocene period (giant jaguar of North America, P. atrox). There are four present-day species in the genus: tiger, P. tigris Linnaeus, 1758; lion, Р. [ео Linnaeus, 1758; leopard, Р. pardus Linnaeus, 1758; and jaguar, P. onca Linnaeus, 1758. These constitute 11.5% of the species of the family. All these forms constitute a group which is morphologically and phylogenetically very close, but views on the affinities between the various species are quite diverse. Some (Pocock, 1929, 1930, and 1941) do not divide the genus into subgenera, or assuming a large number of species in the genus (Haltenorth, 1936 to 1937) place the foregoing species in a single subgenus; other authors (Hemmer, 1966) classify them into two subgenera by placing tiger in a subgenus, treating it as the most primitive form, and the remainder in a special subgenus (Ellerman and Morrison- Scott, 1956 and 1966; see list of synonyms). Most taxonomists consider the lion the most primitive form of the genus and see in the tiger the species closest to lion. It has, however, been suggested (Hemmer, 1960), based on the sum total of morphological and ethological features, that the tiger should be regarded as the most primitive and lion the most specialized form of the genus. In the present work a purely morphological series—lion— tiger—leopard—has been adopted without classifying them into subgenera. The maximum number of species (three) occur in the Old World, one in Africa and Asia (leopard), one in Africa, Asia, and Europe (lion), and one purely Asian (tiger). There is only one species (jaguar) in the New World. Panthera are very valuable game animals and their fur highly prized. They regulate ungulate populations and at places destroy agricultural pests (tiger, wild boar, African leopard, baboon). In some areas they also destroy catile and pose a danger even to man. Russian fauna includes three species: Поп, Р. [ео Linnaeus, 1758; tiger, P. tigris Linnaeus, 1758; and leopard, P. pardus Linnaeus, 1758. These comprise 75% of species of the genus, about 11.5% of species of the family, and about 1.0% of species of Russian fauna. Lion had already become totally extinct in the tenth century. The range covered the southern montane and forest region, at places semidesert and desert parts of the country, and has shrunk markedly in the last decade. These are forest and montane forms, and inhabitants of reed thickets. In view of their small numbers, their economic importance is negligible and rarely adverse. One species (tiger) is totally protected; like the leopard, it is of much interest to zoological gardens. (V.H.) 76 83 LION Panthera leo Linnaeus, 1758 1758. Felis leo Linnaeus. Syst. Nat., 10th ed., vol. 1, p. 41. Constantine, Algeria. 1826. Felis leo persicus. Меуег. Diss. inaug. anatomicomedica de Genera Felium, р. 6. Persia.*> (V.H.) Description Animals of large size (one of the two largest species of the family). General appearance typical of big cats. Body greatly extended, tail length about one-half of body length, and height at sacrum not more than height at shoulders. Head relatively large and heavy, and markedly extended (with a large facial region). Legs strong with broad and powerful paws. Tail with tuft of long (black) hair at end (Fig. 34). Sexual dimorphism very pronounced, much more distinct than in any other species of the family, and very characteristic. Coat of females even, short, fairly stiff, and uniformly dense sandy- or brownish-yellow color without pattern. The difference in color of flanks and abdomen, and outer and inner surfaces of legs, the latter being somewhat lighter, is insignificant and devoid of sharp delineation. Black spot occurs on back of ear. Hair on flanks and upper surface of neck turns up and forward, and elsewhere on the body downward. Lower surface of trunk and inner surface of legs usually with blurred pale spots, darker and more rust-colored than main background, but faint. Frequently such spots are also visible on flanks and even form very faint vertical rows. Black tuft at end of tail somewhat smaller in female than in male. Rarely, rudimentary mane visible in very old females, con- sisting of very short and sparse hair covering a small area on the head and neck. Males and females are similar in color but the former have a large mane of dense hair which, in old animals, reaches 40 cm in length. The mane, although forming a single unit, can be divided into several parts: one segment (chest mane) includes rear of head (commencing from ears), sides of neck both above and below, longer hair on sides of head (whiskers), and a large mass on chest and between forelimbs. Elongated hair (abdominal mane) extends in the form of a band along the abdomen (sometimes along its entire Additional synonyms have been given by В. Pocock (1930 and 1939). ‘лолешоу *N’W Аа 451246 `9С8Т ‘12АЭИ 2215424 02] влащит “UOTT ‘уе "ЗЫ 85 length). Another broad band of long hair covers the underside of the flanks from the rear of the axilla up to the groin region (lateral mane). Hair on the rear of the forelegs from the ulna downward is likewise elongated. It is longest on the ulna and often forms tufts; below the ulna it is shorter and finally disappears. From the neck and nape long hair extends backward, covering the back in the region between the shoulders (shoulder mane) or even slightly farther back. It may cover the upper part of the shoulder region or form a cape along the rear of the scapular region. This cape may extend downward so far that it merges with the abdominal and lateral manes (see Fig. 39). Mane growth commences at the age of six months. Gradually increas- ing in density, size, and luxuriance, the mane is fully grown by the third year but attains maximum growth and шхипапсе in old animals. The mane consists of yellow-colored hair, corresponding to the general color of the trunk, with an admixture of more or fewer black hair, and some of lighter color—white, or very light yellow hair. The latter are more common in the frontal (facial) portion of the male. A large part of the mane, mainly the rear portion, is black and darkens even more with age. The general luxuriance, density, and color of the mane, as well as the extent of its growth (form and area covered) exhibit individual variability. Geographic variability is also significant. The most luxuriant and extensive manes (tresses on flanks and abdomen) occurred in extinct lions of the Cape region (P. /. melanochaitus) and northwestern Africa (Atlas; Р. [. leo). The mane is less developed in other forms, and in some covers only the head, neck, and front of the chest.4 Faint spots are sometimes seen on males, but rarely so; the spots are vague and do not cover large areas of the body. They are seen only on the abdomen and paws and never on the flanks; in rare cases traces of them might be discerned on the latter. The final feature of sexual dimorphism in lion, as in all cats, is manifested in the smaller size and much lighter build of females. Newborns are, as a rule, covered in distinctly circular dark spots, which tend to form vertical rows; in some animals these spots even fuse into distinct vertical stripes. Sometimes newborns are monochromatic and have only faint dark spots on the underside. Usually this skin pattern has gradually disappeared by the age of about six months; more rarely it is preserved for two to three years or even longer. Females have been known to preserve abdominal skin spots throughout their life. *4A particularly luxuriant mane is grown by caged lions. The mane of wild animals is often damaged by their movement through underbrush and bushes. UY 86 The skull is very similar to that of tiger (see below) but usually some- what more depressed and flattened in the frontal region, with a slightly shorter postorbital region. Hence the facial part, compared with the cranial, appears longer and more massive than in tiger. The nasal openings are usually broader but the nasals quite short, not projecting beyond the rear of the maxillae (Fig. 35). The inner cusp of the upper carnassial tooth is very small. The lower profile of the lower jaw bulges and hence when placed on a flat surface the jaw rocks; contrarily, the lower jaw of a tiger is concave and does not rock. These features differentiate the skull of these two species, but the individual variability of tiger skulls is so great that only the structure of the lower jaw constitutes a reliable feature for positive differentiation> (Fig. 36). In general body dimensions and skull the lion is similar to the tiger. The weight of an adult male lion is usually about 175 kg and of a lioness about 125 kg. However, much heavier animals are not rare. Large South African male lions weigh 180 to 226 kg and females 127 to 140 kg. Body length of males about 170 to 180 cm and tail length about 70 to 85 cm. Maximum skull length in males 321 to 401 mm, females 292 to 333 mm; condylobasal length in males 309 to 348 mm, females 263 to 291 шт; and zygomatic width in males 220 to 256 mm, females 188 to 212 mm (data from Roberts, 1959). (V.H.) Systematic Position There is no doubt that the lion is closer to the tiger than to any other cat. Fig. 35. Relationship between nasal and frontal bones in lion (above) and tiger. Sketch by N.N. Kondakov. *5Pocock (1939) thought that this feature was ‘‘absolutely constant’’. However, Haltenorth (1936) established that in roughly 10% of lions (11 skulls out of 113), the lower | profile of the lower jaw was either straight or even concave. 78 87 Fig. 36. Lower jaw and line of its ventral profile in lion (upper line) and tiger. Sketch by N.N. Kondakov. Their identical skull structure bears this out. Both species are evidently the least specialized forms of the genus and lion is usually placed at the head of the series (Haltenorth, 1936-1937 and 1957), although some authors begin the series of big cats with tiger (Pocock, 1939; and others). Recently, a Classification of the genus has been proposed according to which tiger is separated in one subgenus, at the beginning of the series, while jaguar, leopard, and lion are placed in another subgenus with lion at the end of the series, 1.е., the two species are widely separated and represent terminal species of the genus (Hemmer, 1966). In building up this scheme, ecological, ethological, and genetic data have been used, but it is not well substantiated morphologically and is hardly acceptable at present. (V.H.) Geographic Distribution The range (reconstructed) encompasses the open, often desert, expanses of Africa, the Middle East, southeast Europe, and northern India. The range is greatly reduced at present. Geographic Range in the Soviet Union The range (reconstructed) in the Soviet Union is very small and covers only the eastern part of the Trans-Caucasus (Fig. 37). Lions are known from the Trans-Caucasus from the Holocene and became extinct only in the tenth century. In the Trans-Caucasus this cat was not incidental (for example, rare migrations) but rather an important member of the fauna. This is supported by the dispersal of the animal in the past into regions adjoining Trans-Caucasus in the south. These regions formed a single faunal complex, with the Trans-Caucasus representing the northern и | [= Al 0 р ор 5. | | y | ere SEs SS EE CE ey ES сети EEE SEED Re СЕПАЕЕЕЕЕЕхЕЕтИЕЕЕ —s ee == oo ed SD ———E | | | | | | {i \ | Se | | | “ll il all Ns ll (Inns ME Fig. 37. Probable maximum area of distribution of lion, Panthera leo L. in the Caucasus in the Holocene (from N.K. Vereshchagin, 1959, with modifications). V.G. Heptner. border. Commencing evidently in the first centuries of our era, the inhabited area and numbers of the animal steadily decreased in the Trans-Caucasus. However, even in the early Middle Ages lions were evidently fairly common and hunted by shirvanshakhs [native people of Trans-Caucasus]. The total disappearance of the animal from the Soviet Union occurred in the tenth century.” To establish a reasonably precise distribution for the lion is now very difficult. Information in the literature and data available for the Middle Ages and folklore traditions are little indeed. During the period of its greatest extent the range covered all the plains and the foothills of the eastern Trans- Caucasus and extended westward almost to Tbilisi. In the north, bypassing the eastern part of the Caucasus, it formed a large extension from the 6Information about the occurrence of lion, together with tiger, in the mid-1600’s in western Georgia (Mingreliya and Imeretiya) is erroneous (Vereshchagin, 1959). 79 89 Apsheron Peninsula to the mouth of the Samur [river]. Extending then to Araks, the boundary turned east in the form of a narrow projection to Yerevan and a little farther. Its northern boundary then extended in the west into Turkey.?’ From the general nature of the range of lion in the Middle East (see below), intrusions of the animal in the past into the extreme south of Middle Asia, primarily in southern Turkmenia and probably along upper Amu-Darya, cannot be excluded. This is supported by some geographic names involving the term “‘shir’’, i.e., lion, for example Shirabad, a town on the Shirabad-Darya, a tributary of the Amu-Darya, or the Shir-Tepe region on the upper Tedzhen, and others. In fact, there is no direct historic proof of the existence of lion in Middle Asia but it is possible that a study of the source books of the Middle Ages could provide such information. The range of the lion in Persia in antiquity stood quite close to the boundaries of Turkmenia, and the cat evidently prowled even in Afghanistan. Habitat conditions of these regions even in the early twentieth century were favorable for the survival of big predators (tiger, cheetah, wolf, hyaena, and leopard). In the past, and more so in the very remote past, extensive tugais, pistachio savannas, rushes, etc., together with an abundance of kulan, goitered gazelle, wild boar, mountain sheep, and deer constituted an environment which was even more luxurious for the survival of lion. Various views have been expressed about the occurrence of lion, or at least its appearance, in the southern European parts of the USSR. These views are based on the interpretation of the old Russian “‘lyuti zver’ ’’ (fierce animal). The well-known citation, always referred to, are the words of the great Count of Kiev, Vladimir Vsevolodovich Monomakh (1053 to 1125) in his work, Poucheniya Detyam (1117). In describing a hunt in the years of his rule in Turov and Chernigov (1073 to 1094), he wrote that a fierce animal sprang toward his thighs, injuring both him and his horse. The traditional interpretation, suggesting either a wolf or lynx as this ‘fierce animal’’, is evidently unacceptable; these animals could hardly spring at the thighs of a rider, let alone throw a horse. At present some think this “fierce animal’’ was a leopard while others favor lion. The latter interpretation (Charlemagne, 196478, and others) is quite widely accepted. However, there is undoubtedly а good basis for seeing in this “‘fierce animal”’ a tiger (see chapter on tiger below; Heptner, 1969). Thus the appearance 2’The distribution and history of lion in Trans-Caucasus are mainly based on data from N.K. Vereshchagin (1959). 8This work includes a list of the main publications on this subject; also see У. Mavrodin (1965) and V. Sapunov (1965). 80 90 of lion in southern Russian steppes or even the month of the Don, as suggested, cannot be accepted.” In the Pleistocene and Holocene lions were dispersed in Europe far more widely than in the historic past. There is a great deal of information about the finds of leonine remains in southern and central Europe, France, England, Germany, and Poland. Such remains have been reported in the Soviet Union from a large number of sites in the Trans-Caucasus and Crimea as far as Yaroslav and Molotov regions, the Urals, and Krasnoyar.*° At the same time it is extremely difficult to study the distribution of lion in that period since the identification of big cats is difficult and lion is sometimes mistaken for tiger, especially in the case of the extinct tiger- lion (cave lion, P. spelea). This is true, for example, with regard to specimens from Chernigov and others (N.K. Vereshchagin) or the Polish lion, now regarded as P. spelea (Koval’skii, 1959). Nevertheless, some researchers (Hemmer, 1967) consider the big Pleistocene cats lions, and the origin of the species is placed in the Early Pleistocene. Geographic Range outside the Soviet Union The range in Asia and Europe, which in part includes the Trans-Caucasian section of the habitation of lion, and some stages of the disappearance of these animals are as follows. In Europe during the historic period lions inhabited the southern part of the Balkan Peninsula in the north up to at least Thrace and Macedonia and probably even the Danube. Their disap- pearance in Greece occurred at the end of the first century A.D. (Harper, 1945). In Asia Minor the animals were widely dispersed and evidently absent only in the northern part covered by the Pontic mountains (Hemmer, 1967). They disappeared from the western parts of the country [Turkey] evidently much earlier; in the east they survived up to the nineteenth century in regions adjoining Mesopotamia and Syria and disappeared from there only around the middle of that century. In the early 1870’s they were still sighted in the upper reaches of Euphrates (Biledzhik; Danford and Alston, 1877 and °Remains of lions from Ol’via (mouth of the Bug) belong to the fifth to the second century B.C. and are those of animals from a menagerie; some could well be only bones or skins of animals used as rugs. These ‘‘remnants’’ cannot serve as proof of the occurrence of wild lions in Ol’via (Gromova, 1928; Topachevskii, 1956; Vereshchagin, 1959). One must keep in mind that if a distinction between lion and tiger is difficult on the basis of their skulls, it is impossible from the remnants of a skeleton, especially metapodia. It has been suggested that there are no differences between these two species in the postcranial skeleton. At the same time, tiger has never been mentioned in references to Ol’via (see section on tiger here and also Heptner, 1959). (V.H.) For a list of these finds, see I.G. Pidoplichko (1951). 91 1880), but at the end of that century were no longer present. In Mesopotamia and Arabia lions still survived in that period. In the 1850’s they reached the upper courses of the Tigris (Mosul) in Iraq, and in the 1860’s were extremely abundant in reed marshes along the banks of the Tigris and Euphrates, mostly their lower reaches (Blanford, 1876). Lions also occurred in Kurdistan, Armenia, the eastern parts of present-day Turkey, and western Пап (Fig. 38). In Iran they were wide- spread, the Trans-Caucasian range being mainly linked with the Iranian range. In the far remote past they evidently covered a very significant part, if not much, of the country, and reached Teheran in the north, occupy- ing even the Persian upland. In the 1870’s they were sighted only in the south along the western slopes of the Zagros mountains, adjacent to Mesopotamia (east of the Tigris valley), and in the forest regions to the south and southeast of Shiraz (Blanford, 1876). In southern Iran lions were encountered at places, though rarely, even around 900 A.D. (Mekran coast).?! The last of the species was killed in Iran in 1930 (Haltenorth and Trenze, 1956). There is every reason to believe that the “‘king of the jungle”’ prowled even in Afghanistan, at least in the southwestern and southern areas 32 In India the range in the past covered the whole northern sector from Sind in the west to Bengal in the east and from the Narmada River in south (somewhat south of the tropics) to Rampur and Rohilkhand (about 20°30’ NL) in the north (Pocock, 1939). Lions have disappeared throughout this territory except for a small section in the Gir forest in Kathiawar (see Fig. 38) where they have long been protected. These animals are the sole survivors of the Asian lion which, in the past, was not only widely distributed but, in some countries, for example Syria during the Crusades (eleventh to the thirteenth centuries), quite abundant (Usama ibn-Munkyz, 1922). Apart from Europe and the Near Easi, the range covers all of Africa except regions of humid tropical forests in western Africa. By the twentieth century the range had shrunk and lions had become extinct in the southern parts of Africa and in much of the northern mainland. In the 1950’s (see Fig. 38) the animal was present only in tropical Africa from Somalia, Sudan, and Senegal southward to southwestern Africa, Kruger Park in Transvaal, and possibly Zulu- and Swaziland. (V.H.) 3!Personal communication from preparators N.A. Zarudnyi and S.A. Aleksandrov. (V.H.) *°The occurrence of lion in Afghanistan in the past is sometimes doubted. It is inferred here from the nature of the range but direct references to lion in this country are also available. Pocock (1939) assumed the existence of lions in the southern parts of Afghanistan where, according to some data (Harper, 1945), this animal was actually sighted in the middle of the last century. 81 81 SV Fig. 38. Reconstructed species range of lion, Panthera leo L. Broken line denotes roughly the northern and southern boundaries of distribution by the 1950’s. Dot indicates present habitation of lion in India, in Gir, Kathiawar. V.G. у Нершег. Geographic Variation A large number of forms have been described. Of these, eleven are usually recognized: ten in Africa and one in Asian part of the range. Of the African forms, two are totally extinct and one almost so, and preserved only in sanctuaries (see below). There is [was] only one form in the Soviet Union. Persian lion, P. [. persica Meyer, 1826. This lion is wholly similar to the African forms but differs generally in the less swollen tympanic bullae, the much shorter postorbital constriction, and usually by the divided (doubled) infraorbital foramen (Pocock, 1939). The color varies from reddish-brown and highly mottled black to sandy or cinnamon-gray. The mane in males varies in growth and color, being sometimes small but usually dense on the head, neck, and chest, as also on the abdomen and flanks. Ulnar hair is well developed. Usually mane hair is rusty-brown with an admixture of blackish and white hair; it may also be totally golden-brown or contain a goodly admixture of black hair along the top and chest. Ulnar hair and the abdominal and lateral manes are 82 82 93 relatively more dense than the mane ргорег.? The tail tassel is large (Pocock, 1939; Hemmer, 1967; Fig. 39). In overall size the Persian lion corresponds to central African animals. Body length of adult males about 195 cm (two animals) and tail length about 80 to 90 cm (two animals). Maximum skull length in adult males (three) 330 to 340 mm and in females (three) 292 to 302 mm; condylobasal length in males 299 to 313 mm and in females 266 to 277 mm; zygomatic width in males 221 to 231 mm and in females 195 to 209 mm (data for Gir lions; Pocock, 1939). This species used to exist in Trans-Caucasus (up to the tenth century A.D.). Outside the Soviet Union this species used to live in Greece, Asia Minor, Syria, Palestine, the Arabian peninsula, Iraq, Iran, Afghanistan, and Pakistan, and presently lives in the Kathiawar peninsula of India. Evidently lions were not entirely identical throughout the above territory. In Greece and Asia Minor animals with a less developed mane, without abdominal and lateral manes, and without ulnar hair predominated. In Syria and countries on the east coast of the Mediterranean Sea, and possibly the Arabian peninsula, lions of this type also lived, which were very similar and/or identical to the Egyptian lion (P. [. leo Hemmer, 1963).34 There Fig. 39. Main types of manes of Middle East (Persian) lion, Panthera leo persica Meyer according to Hemmer, 1967 (mane hair shown in black only for purposes of visual effect). Sketch by N.N. Kondakov. 33In literature, especially popular literature, the view prevailed that the mane of Persian-Indian lion was very small or totally undeveloped, or almost so. Descriptions of maneless lions of Gujarat are available. This misunderstanding is based on the title of an article published in 1833 by Officer Smi, one of the first to describe Indian lions. Compared with lions of the Atlas mountains and menageries in which the mane is large, the mane of Gujarat (Gir) animals appeared small to him; he used the unfortunate expression **maneless, `` which has come into wide usage (for more details, see Pocock, 1939). 34Вазе on’a study of extensive material, i.e., ancient Greek, Hittite, Aramaic, Urartaean, Assyrian, and Babylonian works of art, H. Hemmer (1963, 1966, and 1967) reconstructed the external appearance of lions of Europe and the Middle East. 94 were all types of transitional forms to the above-described Persian lion, which was the lone or predominant form in Kurdistan, and apparently in the Trans-Caucasus (Hemmer, 1966, and 1967). bY Bd ts The following forms are usually recognized for Africa: 1) Berber Поп, Р. /. leo Linn., 1758—North Africa from Atlas to Egypt (extinct); 2) Senegal lion, P. 1. senegalensis Meyer, 1826—from Senegal to Nigeria; 3) Cameroon lion, Р. [. kamptzi Matschie, 1900—northern Cameroon and the region south of Lake Chad; 4) Congolese lion, P. [. azandica Allen, 1924—northeast Belgium Congo; 5) Masai lion, P. [. massaica Neumann, 1900— Ethiopia, Kenya, and Tanganyika up to Mozambique; 6) Uganda lion, P. /. nyanzae Heller, 1913—Uganda; 7) Katanga lion, P. |. bleyenberghi Lénnberg, 1914—-Katanga, Angola, and Rhodesia; 8) Transvaal lion, P. /. krugeri Roberts, 1929—Transvaal and Orange Free State; 9) Kalahari Поп, P. /. vernayi Roberts, 1948—Kalahari (Botswana), southwest Africa, and southern Angola; 10) Cape Поп, P. 1. melanochaita H. Smith, 1858—Cape Colony to Natal (extinct); and 11) P. 1. hollisteri Allen, 1924—northern bank of Lake Victoria. Most of these forms have been described from extremely variable and insignificant characteristics, and hence their status is dubious. It has been suggested that all African lions are identical in subspecific features. (V.H.) Biology There is no direct biological information about the Trans-Caucasian lion. This lion lived in an environment different than the one prevailing at present and only under those conditions was its existence possible. The country was thinly populated and uncultivated or, in the Middle Ages, cultivated only here and there. At the same time the population of domestic cattle, on which the animal depends for survival at many places even today (Gir and some regions of Africa) was large in the remote past in Trans-Caucasus. Wild ungulates were additionally abundant. Not only large rivers like Kur’ and Araks, but also small streams had rich forests along their banks; dense reed thickets grew along the extension of the Mugansk steppe and at other places in the lowlands; pistachio and juniper forests covered the foothills and there were rich pastures. All these supported a large number of deer, roe deer, wild boar, while in the steppes were large numbers of goitered gazelle and kulan. Huge reserves of ungulates were available to lions as food (cheetah 95 also fed on them). The disappearance of Поп is naturally associated with an increase in human population and a change in environmental conditions, which in turn led to an impoverishment of ungulates in the country; thus, kulan had disappeared from the Trans-Caucasus in the thirteenth century. Indubitably, however, direct killing also played a significant role. Big predators in general are few and hence easily exposed to hunting. Further- more, lions from the time of the ancient kingdoms of Egypt and Mesopotamia represented a favorite object of hunting by emperors, pharaohs, and feudal kings. Lions were also hunted by the shirvans of Trans-Caucasus who, of course, were not the only ones responsible for their being killed (Vereshchagin, 1949). At present, this subspecies is preserved only in the Gir preserve in Kathiawar peninsula (northwest India); the population is very small and rapidly declining in spite of protection. By 1960 the lion population according to official information was 283, but actually there were probably not more than 150 (Ulibrich, 1962); in 1963 they were placed at 285 but by June, 1968 only 162 remained (UCN Bulletin, no. 8, р. 61, 1968). (V.H.) TIGER Panthera tigris Linnaeus, 1758 1758. Felis tigris. Linnaeus. Syst. Nat., 10th ed., vol. 1, p. 41. Bengal. 1815. Felis virgata. Mliger. Abh. К. Akad. Wiss. Вен, р. 98. Mazanderan, northern Iran. 1842. Felis mongolica. Lesson. Nouv. Tabl. Regne Anim., p. 50, Bukhara, Mongolia, Kirghizian steppes. Nomen praeoccupatum. 1844.°° (1845 and 1847). Felis tigris altaicus. Temminck. Fauna Japonica. Mammalia, р. 43. Когеа.36 1858. Tigris striatus. Severtsov. Rev. Mag. Zool., vol. 10, p. 386. Renaming of Felis tigris Linnaeus, 1758. 1867. Tigris regalis Gray. Proc. Zool. Soc. London, p. 263. Renaming of Felis tigris Linnaeus, 1758. “The year of description of this form was originally placed as 1845 (Brongersma, 1935; cited by Mazak, 1965) and later corrected to 1844 (Mazak, 1967). Some specimens from the collection of Temminck (examples in the library of V.G. Heptner) bear the date 1847. (V.H.) >More precisely, Pisihan mountains roughly at 43° NL. This parallel traverses slightly south of Vladivostok, through the extreme northern part of the Korean People’s Democratic Republic and the southern part of northeastern China (Manchuria), i.e., north of the Korean Peninsula. (V.H.) 1868. Felis longipilis. Fitzinger. 512. Ber. Akad. Wiss. Wien, vol. 58, р. 435. Amur region.%’ 1871. Felis tigris var. amurensis. Dode. Proc. Zool. Soc. London, p. 480. Ussuri estuary, Khabarovsk.%8 1904. Felis (Tigris) septentrionalis. Satunin. Zh. Priroda i Okhota, no. 7, p. 5. Twelve versts west of Lenkoran, eastern Trans-Caucasus. 1904. Felis tigris coreensis. Brass. Nutzbare Tiere Ostasiens, p. 4. North Korea. 1915. Tigris mikadot. Satunin. Zh. Nasha Okhota, по. 7, р. 17. Korea. 1916. Felis tigris lecoqui. Schwarz. Zool. Anz., vol. 47, p. 351. Lobnor, Kurla. 1916. Felis tigris trabata. Schwarz. Ibid., vol. 47, р. 353. Ш valley south of [Lake] Balkhash. 1925. Felis tigris mandshurica. Baikov. Manchurian Tiger, p. 3, Harbin, northeastern China (Manchuria). 1929. Tigris mandjurica. Baikov. Zh. Okhotnik, no. 4, p. 16. Substitute for mandshurica Baikoff,* 1925. 1929. T. koraiensis (micadoi) Baikoff (ex Sat.). Baikov. Zh. Okhotnik, no. 4, p. 16. Northeastern China (Manchuria). Substitute for koreensis Brass, 1904 and mikadoi Satunin, 1915. 1935. Tigris tigris virgata Matschie (1897 ex Illiger, 1804 to 1811). Ognev. Zveri SSSR, vol. 3, p. 273. Substitute for virgata Mliger, 1815. (У.Н:) Diagnosis Color reddish-rusty or rusty-yellow with narrow black transverse stripes. Body length not less than 150 cm, condylobasal length of skull 250 mm, zygomatic width 180 mm, and length of upper carnassial tooth over 26 mm. (V.H.) Description*? Size very large; largest species of the family. In general build the tiger is wholly typical of cats and has an extended 3’Locality fixed by S.I. Ognev (1935) and adopted by other researchers. S.U. Stroganov (1962), based on the fact that Fitzinger first indicated *‘Middle Asia’’, placed Lobnor Basin as the type locality. 38According to S.U. Stroganov (1962). This work also includes a discussion of the names of Russian tigers since 1868 from a nomenclatural point of view. *Spelling of author’s name in taxonomic divisions is sometimes at variance with spelling in text and bibliography since the Israeli orthography has been followed in this translation. ЭТп the ‘‘Description’’ and ‘‘Geographic Variation’’ sections the works of $.1. Ognev (1935), S.U. Stroganov (1962), and V. Mazak (1965 and 1967) have been incorporated, in addition to works cited in the text. The characteristics given under *‘Description’’ have been supplemented and elaborated in ‘Geographic Variation’’. ‘лолешоу ‘КУ Aq 451246 “1 $м8и плащи Г ‘19817, ‘Op ‘314 Sg 98 86 Fig. 41. Amur tiger, Panthera tigris altaica Тетт. Moscow Zoological Garden. Photograph by A.P. Zhandarmov. supple body standing on rather short legs with a fairly long tail (Fig. 40). The front portion of the body is better developed than the rear and the animal stands higher at the shoulders than at the sacrum. This gives the impression of some heaviness but simultaneously of immense strength, which is emphasized by the powerful legs with large and broad front paws, heavy and invariably slightly drooping head, high withers, and prominent strong musculature, especially in the anterior part of the body, primarily in the form of powerful shoulders. The animal is quite slow on its feet and even its more rapid movements, leaps for example, appear leisurely. The same impression of quiet strength comes across even when the tiger is lying down. The general appearance of tiger is that of a huge physical force and quiet confidence, combined with a rather heavy grace (Fig. 41). Head with markedly projecting facial portion and bulging forehead. Ears short with a rounded, blunt apex and wide-set. Vibrissae set in four or five rows, white, sometimes brownish at the base, extremely resilient, up to 165 87 99 mm long, and up to 1.5 mm thick. Pupil round and iris yellow. Claws very powerful, sharply curved, and a pale waxy color. Their length along the outer arc reaches 80 or 100 mm and height of base up to 40 mm. Summer coat rather short, quite sparse, of uniform length throughout the body, with long hair present only on cheeks. Some long hair is also seen along the upper part of the neck. Coat quite coarse. In southern forms living outside the Soviet Union there is no difference between the winter and summer coats, or very little. Among tigers of Turkestan and the Caucasus, but especially among those of the Soviet Far East, coat differences are very sharp: winter coat is denser and longer, softer and silky, and might even appear shaggy on the trunk of animals with an especially rich coat. Hair is markedly longer on the head (the ears project insignificantly); whiskers and hair on the occiput and along the top of the neck also greatly elongated. Compared with other Soviet cats, the pelage of the tiger is of moderate thickness, coarse, and sparse. Back with 2,500 hairs per 1.0 cm and abdomen with 600; 1.4 underfur hairs occur for every 1.0 guard hairs (an extreme case among Soviet cats). Underfur very slightly curled and coarse. Exclusively black hairs (of black stripes) differ greatly from yellow hairs, 1.е., those devoid of black pigment. Average length of yellow hair 27 mm and thickness 105 microns and for black hair 48 mm and 88 microns respectively. Length of yellow guard hair (categories I to ГУ) 31, 29, 25, and 25 mm and thickness 85, 62, 49, and 37 microns; length of black guard hair 40, 38, 33, and 31 mm and thickness 87, 63, 47, and 34 microns. Yellow underfur 24 mm long and 30 microns thick and black 23 mm and 30 microns respectively (winter skin of Amur tiger; B.F. Tserevitinov; also see “Geographic Variation’’ below). Main background color bright reddish-rust or ocherous-rust. This shade is brightest on the back and less vivid on the flanks and outer side of the legs. Lower part of head, neck, midportion of chest, abdomen, and inner side of legs white or with light yellowish tinge. Transition of color from upper to lower surface gradual but distinct. Main color of tail on dorsal surface same as on trunk, but toward the end slightly lighter, and may even become white; underside of tail white or very light ocher. Against this main background color an extremely complex pattern of dark (pure black or brownish) stripes is distinct. These stripes vary in size, form, and disposition in different parts on the body; vividness and sharpness of outline also vary. There are no annular spots on the coat. The following disposition of dark stripes is typical: transverse (vertical) position throughout the body and along the neck; on the trunk some descend low while others terminate above, or even extend onto the abdomen. All stripes with sharp lower ends, sometimes bifurcate, or divide and fuse again 88 100 toward the end. Other transverse stripes commence between these tapered ends and descend onto the abdomen or encircle it. Those commencing on top of the neck terminate on the neck or run onto the chest; those from the withers descend along or onto the rear of the shoulder blades. Stripes are usually more densely distributed on the posterior half of the trunk than on the anterior. Anteriorly, however, the color is usually more intense—more or less pure black—and not uncommonly a brownish shade posteriorly; they may even be brownish on the thigh. Stripes commencing on the sacrum run onto the groins or to the rear of the thigh, or terminate midway on the thigh. Some, having reached the upper part of the coat, take an oblique direction, incline forward, and run onto the inner surface of the legs. The outer and inner surfaces of the hind legs up to the calcaneal joint are covered with dark transverse (horizontal) stripes, which are short or terminate in a semicircle or even a ring. Foot dirty gray below. Front (outer) surface of forelegs devoid of black stripes, while inner surface carries short, intermittent black transverse stripes, their ends sometimes extending onto rear (outer) surface of legs. Lower ends of stripes on shoulders and in scapular region may descend to the ulna. Tail invariably with a black tip and for much of its length covered with transverse bands forming compact rings. Such rings are absent, however, at the base of the tail; dark markings occur along the top of the tail in the form of angles with apices turned forward. There are usually 10 dark transverse stripes on the tail but these may number 8, 9, or 11. The pattern on the head is complex (Fig. 42). Dorsal side of muzzle, commencing from nostrils and up to region between the eyes and slightly posterior, rusty-colored and without dark pattern. Region from the corner of the eye to upper row of vibrissae similarly colored. This coloration usually extends in the form of a small wedge under the eye to its outer corner. A narrow white field runs directly along the lower eyelid. Region of lower nostrils and vibrissae, chin, and lower jaw white except for black spots on lower lip in the corner of the mouth. Sinciput and occiput with complex and variable pattern of short transverse black stripes, often irregular in form; there are usually four, five, or six such stripes. Sometimes some or all are interrupted in the middle; sometimes a longitudinal stripe occurs between them and transverse stripes are set off from it. The last pair of transverse stripes sometimes forms an angle.*? 40Тре head pattern consisting of a longitudinal and three transverse stripes (see Fig. 91) forms a figure which is extremely similar to the Chinese hieroglyph ‘‘уап”” or “‘уап- da*’ which means *‘great ruler’’. Among the Chinese and Koreans and some smaller tribes of the Far East, tigers were associated with numerous superstitions and animals with the sign ‘‘van’’ considered sacred and treated respectfully (Baikov, 1925). 101 Fig. 42. Amur tiger, Panthera tigris altaica Temm. Moscow Zoological Garden, 1963. Photograph by У.Е. Prisyazhnyuk. Often a long stripe begins behind the ear and descends in the form of an arc onto the neck, often reaching the throat. The sides of the head and the whiskers around the white field are covered with some black stripes of angular or bracket shapes. At their top they commence under the ear or before it. Sometimes one of the transverse bands of the sincipital pattern also extends down along the cheeks and flanks, reaching the corner of the lower jaw or even descending onto the throat. The front surface of the ear is covered with white hair while the rear is invariably pure black with a large bright white spot in the upper half.*! The above description of tiger coloration is highly schematic (see also ‘““Geographic Variation’’) and undergoes a good deal of variation, primarily individual; color shade as well as pattern undergo changes. The general color of the coat is sometimes quite vivid and bright rusty and sometimes a dull yellow. The downward extension of the rusty field also varies. So too do the white fields on the underbody, rising to various degrees on the flanks “lThis spot, developed in some other large (leopard) as well as small (Amur [= leopard] cat) cats, is of significance as a signal. 90 102 and neck. The coloration of the pattern itself likewise varies. The black stripes on the trunk may be brown-toned while those on the thighs and rear of the paws, usually somewhat brownish, may be more vivid. The black pattern is subject to considerable variability. Stripes may be wide, narrow, continuous or bifurcate at the end, or with a light-colored interruption, shorter or longer, several or many, the skin densely or sparsely covered with them, and so on. Stripes on the flanks usually broaden in the middle of the back and, converging with the stripes from the opposite side, merge or alternate with them; sometimes a black longitudinal band forms on the back, often extending the entire length of the trunk. Sincipital pattern and that in general on the head, side of the muzzle, and around the eyes varies notably. Pattern of tail likewise is highly variable. In addition to‘distinct and well-demarcated closed rings throughout the tail one also sees tails with only a black tip and some distinct distal rings, or tails with paired dark rings and yellow bands. More often the front rings are not distinct—neither completely covering the tail, irregular in form, and without a distinguishable pattern. By and large these ‘‘rings’’ are angular bands distorted to various degrees, but with the apex turned forward. Nevertheless, whatever the variation, ten dark fields are invariably distinguishable. The background color of the tail may be white throughout its length or for most of it. Tigers showing sharp deviations are known. Some have a reduced number of black stripes in the front half of the body. In extreme cases the scapular region and chest are almost devoid of a black pattern, while in the rear portion of the body stripes may be normally developed. Melanism is known, as is albinism (some faded traces of stripes are usually discernible), and so are referred to as white tigers. In the latter the rust-colored pigment is reduced but the black pattern fully developed. These mutations have not been seen among Soviet animals; a reduction in number of stripes has been observed very rarely. Seasonal color variations in northern, 1.е., Soviet, tigers are significant. The main background coat color in the winter, compared with the summer coat, is less bright and rusty, in other words more ocherous. As a result of the greater length of the pelage, the stripes appear broader and have less sharp outlines. Sexual dimorphism in color is not apparent. Whiskers in the winter coat are not so well developed in females as in old males. Age-related color changes are absent; young cubs have the same striped coat as adults but the main shade of their coat is less vivid and more ocherous. Moreover, the stripes in cubs are distributed closer together, are wider, and their outlines less sharp, due partly to the relatively long hair and furriness of the coat. A black line on the back is usually present. 103 Geographic variation in color and pelage type is clearly expressed and reflected in the intensity of the rust coloration; the extent of development of the stripes (their number, density, width, and length), intensity of black coloration, size of the white field on the underside of the body, the black pattern on the tail, and other details of color also undergo change to some extent or another. The degree of seasonal dimorphism in length and richness of the coat also undergoes geographic changes. The geographic color changes follow Gloger’s rule (more details later). Albinism and melanism are known among tigers living in the wild. Under certain light conditions traces of dark stripes are usually seen among albinos. White tigers, present mainly in America, are those raised in zoological gardens from a lineage of animals in which the yellow pigment has failed to develop but the black pattern has remained. These animals owe their origin to the tiger indigenous to India. Teats; two pairs, inguinal. Diploid number of chromosomes, 38. The skull (Fig. 43) is wholly typical of the genus and characterized by its large size. In this respect, as also in almost all others, it is similar to the skull of lion, from which it differs only in structural features of the lower jaw and relative length of nasals described above (see description of lion).4? The skull of the tiger is heavy and massive with a large and very powerful facial region, and very broad in the region of the canines. The cranium is elongated and powerfully compressed with well-developed crests, especially the sagittal. General skull profile bulges slightly, the highest point being the interorbital region. Orbits are relatively small, oval in form, extend to the rear, and open broadly behind. Tympanic bullae are relatively small, thick-walled, and flattened. Mastoid process broad, massive, and conical in form. Lower jaw heavy, with a broad coronoid process. Lower profile of jaw somewhat concave (see Fig. 36). Upper canine unusually well developed, broad at the base, and long (to 65—66 and even 70 mm along front curvature). Upper molar very small and sometimes absent (lost). Upper carnassial tooth very large, with well-developed antero-internal cusp displaying a small cone; quite often an additional tiny cone develops on the anterior outer edge of the tooth. Sexual dimorphism of the skull is evident only in the slightly reduced overall size in females, and in old males, intense development of the crest, greater compression of the postorbital region, and also of the brain case itself. “Pocock (1939) pointed out that differentiation of the skulls of lion and tiger is not always easy when a large number of both are to be identified. It is also known that there are no differences in the post-cranial skeleton of the two species. ie SSN 89 Fig. 43. Skull of Amur tiger, Panthera tigris altaica Temm. №. $ 2088, collection of Zoological Museum, Moscow University. Ussuri region. Sketch by N.N. Kondakov. 91 105 Fig. 44. Age-related changes in general features and relative size of skull of tiger. Left—skull of an old male, No. $ 2088, Ussuri region, Bogomolov, with greatest skull length of 360 mm. Right—skull of young animal, No. $ 58977, Ussuri region, between Khor and Iman Rivers, winter of 1955-1956, with greatest skull length of 220 mm. Collection of Zoological Museum, Moscow University. Sketch by N.N. Kondakov. Age-related changes in the skull are very significant (Fig. 44). The skull of young animals is rounded with relatively poorly developed and less massive facial region, a more bulging and relatively large brain case, and zygomatic arches which are weaker, not so angular, and bulge less. With age the skull elongates; the facial region develops at the expense of the cranial part; and the skull becomes heavier, thicker-walled, and uneven. On the whole the range of age-related skull changes is great, the greatest in the family (as in lion). Geographic variability in skull structure is small and pertains only to some differences in overall size. 92 106 The tiger is the largest species of the family, its body size and weight being more than that of the lion. At the same time the largest skulls of lion are only slightly bigger than the largest skulls of tiger.4° The lion evidently has а somewhat larger head and a shorter body, which is generally less massive.** In size and weight the tigress is much smaller than the male. Differences in old animals may reach 15 to 20% in body length and up to 50% in weight. By and large, however, size in these animals is extremely variable; this variability resulting not only from the common individual variations, but also from the fact that tigers live long, continue to grow for a long period (males almost throughout their life), and under certain conditions may become extremely plump. The absolute body and skull size and weight of Soviet tigers, which provide an idea of the species, are given under ‘“‘Geographic Variation’’. The geographic variability in body and skull size is distinctly manifested and follows Bergmann’s rule, but not rigidly. The dimensions of some internal organs based on the measurement of three wild adult tigresses follow (No. S 34885, Ussuri region, Zoological Museum of Moscow University, body length 172 cm, weight 145 kg; No. 197, Pyandzha, Tadzhikistan, body length 165 cm, weight 135 kg; and No. 203, Kyzyl Su, Tadzhikistan, body length 165 cm, weight 97 kg; data of Chernyshev, 1958, with modifications, and V.G. Heptner): weight of heart and heart index 770 g/5.28; 570 g/4.22; and 970 g/10.00 (V.H.); weight of lungs 850, 1,480, and 2,070 5; weight of liver 1,720, 1,955 and 2,179 g; weight of spleen 265, 290, and 250 g; weight of alimentary canal 2,500 and 2,700 g; weight of stomach 795 and 910 g; weight of kidneys 990 and 1,160 g; total length of intestines and ratio to body length 742 (1: 4.31), 640 (1 :3.88), and 710 cm (1: 4.30); length of small intestine 542 cm (87.5% of total length of intestines), 550 (85.9%), and 660 cm (92.9%); length of large intestine 100 (13.5%), 90 (13.4%), and 50 cm (5.6%); and length of appendix 6.0 cm (0.8%), 5.0 cm (0.7%), and 10.5 cm (1.5%). Cross section of heart 118 mm and length 132 mm (specimens of the Zoological Museum of Moscow University). (V.H.) Systematic Position In spite of considerable differences in the general appearance between lion and tiger, these two species should be regarded as very close to each other. The differences in their skulls, as shown above, are very insignificant and “The maximum skull length of tiger is evidently equal to 385 mm (V.G. Heptner; see later) or possibly 400 mm (Baikov, 1925). Among lions the maximum skull length may reach 390 and even 400 mm (Roberts, 1962), the world record being 419 mm (Best et al., 1962). “4It has been shown (Best et al., 1962) that in India, the lion is ‘‘inferior’’ to the tiger. 93 Fig. 45. Amur tigress aged 3.5 to 4.0 years with relatively sharp and broad stripes in anterior body and narrower and closer set stripes in rear. Body length 159 cm, tail length 88 cm, and height at withers 79 cm. December 11, 1961, Suchan River, southern Ussuri region. Photograph by V.K. Abramov. often inconstant; the same is true of differences in the postcranial skeleton. While some subspecies of lion and tiger differ greatly, at the same time resemblances, for example between tigers and lions of Asia, are quite significant. This fact has provided a basis for some zoologists to place the origin of these cats in southwest Asia. On the whole the differences between these two species, taken in toto, are less in craniological features than between other proximate species of cats. Most systematists place these two species in immediate proximity in taxonomic tables. Yet despite the foregoing, Hemmer (1966) has expressed the view that tiger is a distinctive monotypic subgenus and stands in contrast to all other big cats, which constitute a different subgenus. Furthermore the lion consti- tutes a species distant from tiger but most closely linked with leopard, and through it with jaguar. In such a sequence the ethological peculiarities of the species, as also the much easier hybridization of lion with leopard and jaguar than with tiger play an important role. There is no,justification at present to adopt this scheme; it runs contrary to morphology. (V.H.) Geographic Distribution Tigers live in forests and reed jungles in southern, southeastern, and Middle Asia, and parts of the Near East and Central Asia. 108 Geographic Range т ше Soviet Union The range in the Soviet Union (reconstructed) constitutes the northern rim of distribution of the species. Within the Soviet Union the range is divided into three independent sections— Caucasian (including the European part of the USSR), Middle Asian (including Kazakhstan and western Siberia), and Far Eastern (including Trans-Baikal). These areas join in the south out- side the USSR. The Middle Asian, and part of the Far Eastern range, actually consists of isolated sections or pockets (Fig. 46). In view of the already mentioned peripheral nature of the range in the Soviet Union, two parts have been identified, or are being identified, in each of the three habitations of tiger in this country; these are territories of permanent habitation and breeding, and areas where the animals are simply intruders. Usually the region of intrusion adjoins that of a somewhat wider zone of permanent habitation. In relation to the entire range of the species, the area of transgression in different parts of the periphery cannot be very large. However, given the relatively small sections of the range falling within the Soviet Union, this area is quite significant. At places the area of transgression could conceivably exceed the area of permanent habitation. One must also keep in mind that the separation of these two zones is not always possible; thus the true boundary of permanent habitation of the species can hardly be delimited when discussing the past or the period of disappearance of the animal from any part of the range when its normal population and life pattern were disturbed. Tigers are highly mobile and intrusions into other places are frequent. They may be seen for several years almost regularly in a certain area, or for some period become residents and even breed, then for some years disappear altogether, or occur as rare intruders. Sometimes the boundaries shift in a discernible direction. Finally, in some wide regions in the past the animals settled, but later under the impact of extinction appeared as intruders. Caucasus. In the nineteenth century (Fig. 47), the permanent habitation and evidently also breeding grounds of the tiger covered only the extreme southeastern part of the country—Talysh and Lenkoran lowlands. Here tigers were sighted in hilly forests up to the northern extremity of the mountains and in lowland forests. They were encountered most often in the lowland forests of Prishib. Here their population was quite large until the end of the last century and even early in the present century, although significant fluctuations were observed at different periods. A rapid decrease in the number of animals was recorded for the early years of the present century and even the end of the last. The animals stopped breeding and were found only as regular intruders from Iran that remained in the Caucasus for quite some time. This was primarily due to direct ‘лэщ4энН “OA `5э8\/ PIPPI, OY} ит вез илэцуиаоз рие зите4 иезеопео-$10) UI эоцэр!э1 a[quqoid—c¢ ‘59991221 э|4еНэ1 $591 —{ ‘suoisnajut Ав ‘ое — © ‘suoisnyul зиэпБэд] $591 JO 5101891 Jo sauepunog — < ‘uOIsNgUI теп8э1 рие поцеущеч зиэцецио4 Jo suoIdar Jo зэперипод4— | ‘(SOLINJUDD ц}1э^1 01 Ч3иэ}) $э8\/ эр! 94) и! ээцэллийэо Jo $2и$ э1ае9024 рие Аицизо yanuany Apes oy) UL рие циэээщи dy} JO PUD эц зе чош[] 1э1^0$ oY) и! Jay JO э8ие1 рэопдзиооэу “OF "Зы | | | |} =] Yo ar co 4) “oy р y\ ¥ 110 persecution by the expanding Russian populace, changes in topography (deforestation and cultivation of the land), and extinction of wild boar— the main prey of tiger. In the last century, and somewhat even before, when Talysh was a permanent habitation for tiger, the animals moved out from there into different parts of the Caucasus, mostly the plains of the eastern Trans- Caucasus. Talysh later became the site of intrusion of tiger into Russia from Iran. This is because Talysh lies in the westernmost extremity and is covered with the forested landscape of the Cis-Caspian province of Iran that is connected to Elburz (Hyrcania). Evidently only a few tiger intrusions into 95 the Caucasus were not linked with Talysh, occurring instead through the more western parts of northwestern Iran (Zangezur range at Araks in the region of Megra, Armenia; Ararat depression; probably, Akhaltsikha). From 5 00am aa] У Ш J, © (| | \ i bey, г ae ——— aw, ran to Ar, (e) Ye. 29 ‘4 35 Fig. 47. Northern boundary of region of permanent range of tiger in the Caucasus in the nineteenth century (1); individual sites (2); and region of intrusion in the eighteenth, nineteenth, and twentieth centuries (3). V.G. Heptner. 96 111 Talysh the animals most often intruded into the nearest areas of Trans- Caucasus steppe (Belyasuvar in Mugansk steppe, Sal’yana, and others) and also penetrated significantly farther, to Tbilisi, reaching there evidently along the Kur’ valley (1735, 1820, and 1922). Tiger intrusions from Iran into Talysh have become quite rare in the last decade. Reports of such came in the 1950's and 1960’s but the last (to 1969) was recorded in 1966 (Lenkoran low- land).4° The appearance of stray animals even now is not excluded, but its possibility is very remote since the population is rapidly declining in Hyrcania. By the middle of the twentieth century tiger could be considered extinct in the Caucasus.*° In the eighteenth century and the first half of the nineteenth tiger transgressions beyond Talysh were far more numerous and covered the north, east and west more extensively. In the north the animals reached Shemakha (foothills) and Baku and even Derbent; in the east they were sighted in Armenia and Georgia proper (in the east—region of Tbilisi, 1820 and 1835); and in the western Trans-Caucasus in Kvirila and Rioni basins (Imeretiya and Mingreliya), including the southern slopes of the Great Caucasus [range]. The northwesternmost section of the range boundary of the species evidently traversed the latter (Brandt, 1856).*7 The boundaries of permanent hab- itation and region of intrusions are not known. It is quite clear, however, that Armenia and western Trans-Caucasus, like Tbilisi, constituted regions of intrusions. At the same time, probably in the eighteenth century, the animals lived permanently not only in Talysh and Lenkoran lowlands but also along tugais, rushes, and lower reaches of the Kur’ and Araks Rivers. There is justification for believing that in the Middle Ages, especially in the tenth to the twelfth centuries, perhaps even later, tigers lived perma- nently at suitable sites throughout the plains of eastern Trans-Caucasus and, possibly, at some places in the foothills of the Great and Little Caucasus (Fig. 48). The natural environment in these regions was far more favorable for tiger in the Middle Ages than in the nineteenth century. Humans were few, water sources full, tugais and all types of trees abundant along rivers and Some additional information about tiger intrusions into the Caucasus and other parts of the range may also be found in the section ‘‘Biology’’ (*‘Population’’ and ‘‘Seasonal Migrations and Transgressions’’). 46Brandt, 1956; Satunin, 1903 and 1915; Dinnik, 1914; Smirnov, 1922; Ognev, 1935; Vereshchagin, 1947 and 1959; Sludskii, 1953a and 1966. 47Sometimes the data of Giildenstaedt (1787 and 1791), on which Brandt relied, pertain to leopard; these are hardly reliable data. References to tiger in western Trans-Caucasus are also found in other older works (Sharden, Gamba, Nordmann, and others; Brandt, 1856), even for the mountains south of Akhaltsikha in the nineteeth century. 112 lakes and, at places, even along the sea coast; wild boar, deer, roe deer, kulan and goitered gazelle were numerous (see section on lion). Along the Caspian coast the animals evidently penetrated far north. There is every justification for believing that the “‘fierce animal’’ of Russian literature in the Middle Ages was not the wolf, leopard, or a lion, as presumed by many researchers, but the tiger (Heptner, 1969). It may therefore be assumed that tiger occupied even the northern Cis-Caucasian plains; not only the Cis-Caspian rushes far in the north but also Terek, Kuban, and Azov coasts. It colonized even the estuary of the Don and the southern Russian steppe, and probably even the forest steppe (Chernigov principality). Today one must only speculate on the actual outlines of the range and the nature of occurrence of the animal in this extensive expanse north of the Great Caucasus. It is highly probable, however, that the animal yr 4 AEE === Fil cl Fig. 48. Probable range of tiger in the northwestern section of the range in the Middle Ages (tenth to twelfth centuries). 3 97 was a permanent inhabitant, at least in the northern Cis-Caucasus (for more 98 details, see Heptner, 1969). Middle Asian area of occurrence. Within the Soviet Union this region is divided into several independent sections (Fig. 49). In southwestern Turkmenia tigers in the past evidently lived permanently along the Atrek [River] as far as the sea (Chikishlyar) and along its tributaries, Sumbar and Chandyr, in any case along their lower reaches. Insofar as the rest of Kopet-Dag within the Soviet Union is concerned, tigers were encountered there rather often but only as intruders. Tigers lived in southwestern Turkmenia during the last century, but evidently only up to the 1880’s or probably 1890’s. Their disappearance was directly linked with their persecution, mainly as a result of Russian colonization, and also changes in their natural habitat, i.e., the disappearance of rushes along Sumbar and Chandyr and later even along Atrek, and a reduction in population of wild boar. ; In regions very close to the Atrek basin and along Gorgan in Iran, tigers survived well and at places (Talau) lived even until the middle of the present century. From there they penetrated, and continue to do so, into the Soviet Union. Transgressions occur once every several years, mainly into the western and southwestern parts of Kopet-Dag. The animals have been caught on the Atrek, especially at Kyzyl-Atrek and Chat, where four tigers were killed separately in April, 1930 (V.G. Heptner)—at Kara-Kara and at Kaine-Kasyr on the Sumbar, at Nukhur, and even at Kyzyl-Arvat, in the environs of Ashkhabad in the northern foothills. An intrusion is on record into the Great Balkhan, i.e., through the vast expanses of the 4езен. In the 1930’s intrusions were rarer than in the 1920’s and earlier, and in the 1940’s even rarer than in the 1930’s and so on. In the eighteenth century tigers were reported on Cheleken Island (V.G. Heptner) and near Balkhan Gulf (Brandt, 1856, according to Gmelin). The habitation of tiger in southwestern Turkmenia is directly associated with the habitation of the animal along Elburz [range]. The former represents the northeastern extremity of the range, in the same way that Talysh represents the northwestern extremity. Further east, in the expanse between Ashkhabad and Tedzhen, tigers were evidently absent. This region was altogether unsuitable for these animals (Takyr zone at the foot of the Kopet-Dag range) and there are no data even about transgressions. In Tedzhen tigers were reported from Puli Khatun (35° 48The last tiger (to 1969) in Kopet-Dag was killed on January 10, 1954 on the Sumbar around Kaine-Kasyr right on the Iranian border (Shukurov, 1958; in this work data are also available on other finds in Kopet-Dag). It is suggested that some confusion with leopard may have occurred in the reference to the Great Balkhan. However, tigers move long distances across the desert (see below). 114 ОЕ’ ‘ll lon 120 0 120 240360km (ee Se eS | 97 Fig. 49. Middle Asian area of occurrence of tiger in the Soviet Union. 1—boundary of permanent habitation and regular short intrusions in the nineteenth century; 2—sites of stray finds at the end of the eighteenth, nineteenth, and partly twentieth centuries (most intrusions in the twentieth century were in the region of former permanent habitation), 3 — migrations from Amu-Darya delta into Syr- Darya and vice versa, and from Chu River into Semirech’e; 4— not wholly reliable information about Zeravshan and unconfirmed limit of occurrence along Sary Su. V.G. Heptner. 55") up to the river mouth (Zarudnyi, 1890 and 1891). They were undoubtedly encountered even beyond since they bred in Afghanistan along the upper reaches of Hari-Rud (i.e., Tedzhen) at Herat. Along the tugais and reed jungles in the lower reaches of the river at Sarakhs and somewhat higher, the animals undoubtedly lived permanently; they were probably only regular visitors in the much higher levels where large tugais and rushes could not have been present because of the very nature of the river valley (it cuts through the Gyaz’-Gyadyk mountains). 1S Around Tedzhen tigers were common in the 1880’s (especially around Karry-Bend railway station) and were encountered even in the 1890’s. Destruction of forests and ruthless killing led to their total disappearance early in the present century. There is no information for the present century about even intrusions at Tedzhen. The Tedzhen section of tiger habitat was evidently connected with the animal’s occurrence in northeastern Iran, as well as the Afghan section of the range. In the past tigers were extensively distributed and at places very common around Murgab and the state boundary to Mara (Merv) in the north. They were also sighted along the Kushka from Chil’dukhtar (south of Kushka town) up to its confluence with the Murgab. Evidently they also occurred on the Kashan, which enters the Murgab above Kushka. Along the Kushka tigers were seen mostly as intruders though quite regularly, and could have existed as permanent inhabitants in some periods. The disappearance of tigers on the Murgab and Kushka is associated with the destruction of the tugais and cultivation of the valley (cotton), but more so with ruthless persecution, even in the pre-Revolutionary period, especially by hunting parties of the army and some military sportsmen-officers. The animal had already disappeared long before in the north, but persisted in the south until the 1880’s and even 1890’s. On the upper Kushka it was fairly common at that time (1880’s). By the early present century, tigers were practically extinct throughout the Murgab basin and there is no reliable information even about their intrusion into the upper Murgab and Kushka. The possibility of intrusions is not excluded even in the 1920’s since the animal was still sighted though rarely, in adjacent parts of Afghanistan. Even before the middle of the present century, however, they had disappeared there.” The Amu-Darya basin represents a more or less isolated area of occurrence of tiger. The animals occurred from the lower reaches of Pyandzh as far as the Aral Sea. Along this entire stretch two main isolated sections of permanent habitation of tiger occur—northern and southern. In the intervening region stray animals occurred and, if they stayed there at all, residence was brief. Tigers did not flourish, or only rarely flourished, or lived there permanently so long ago that no precise information is available about their prior occurrence. The northern section covers the whole delta of the Amu-Darya where tigers lived in the area northward up to Nukus and Khodzheila. Formerly, tigers were encountered in the tugais along the Amu-Darya, and significantly According to data of Nikol’skii, 1886; Radde and Walter, 1889; Eichison, 1889; Zarudnyi, 1890 and 1891; Silant’ev, 1898; Satunin, 1905; Masal’skii, 1913; Bil’kevich, 1918 and 1924; Flerov, 1932; Leptev, 1934; Heptner, 1949 and 1956; Sludskii, 1953 and 1966; Shukurov, 1958: and others; and data of V.G. Heptner. 99 116 farther, not only at Khivinsk oasis but also in the Dargansk Ata (40°30') and Dayakh-tyn (40°05') tugais. Throughout this region, particularly in the above two southern tugais, tigers were encountered even at the end of the last century and in the early present century, and were found in significant numbers in the delta. Destruction of the tugais, extinction of wild boar, and tiger hunting led to a rapid drop in numbers and a reduction in range. By the 1920’s tigers were sighted only in the delta and even in the 1930’s only around Khodzheila and Nukus. The last of the animals at Nukus was killed in 1938. In the 1940’s they were seen only in the lower regions of the delta _ (in the region on Kungrad, Karauzyak, Muinak, and Kegeili) where some 12 to 15 animals were counted in 1942. In spite of a ban on their hunting, their numbers had declined to about five or six in 1947, and these had disappeared by 1950.2 The southern section of occurrence of the tiger along Amu-Darya includes the Pyandzh valley and the uppermost reaches of the Amu-Darya and its right tributaries. In the east the tiger reached Chubek (south of Kuiyab) and the western foothills of the Darvaza range. It lived permanently among the tugais and reed jungles of the Pyandzh and Amu-Darya as far as Termez, and probably lower. It lived permanently also among the tugais of the Yakhsu, Kyzylsu, Vakhsh, Kafirnigan, Surkhan-Darya, and Shirabad. The animal ascended very high along them—as far as the Gissar valley (upper reaches of Kafirnigan) to the upper reaches of the Surkhan-Darya. The region of permanent habitation and breeding of tigers was connected predominantly with the lower reaches of the above rivers; in the higher parts of their valleys the animals were mostly intruders. This region was the richest in tigers in Middle Asia. This was not only because tigers multiplied undisturbed under the extremely favorable condi- tions in the forests and the unusual abundance of wild boar and deer, but also because their numbers were invariably supplemented by migrants entering the region from Afghanistan.*! This was the situation prevailing According to some data the last of the animals was killed in 1947 and information about killing of tigers in the delta in 1950 and 1951 is regarded as dubious (Sludskii, 1953). However, information is available concerning sightings of lone animals in the delta in 1955 (February), in 1963 (December: Kegeili region), and in 1965 (September; Mambetzhumaev and Palvanniyazov, 1968). Finally, in mid-June, 1968 one tiger, evidently not old, was encountered on the bank of the Amu-Darya 25 km above Nukus. Probably, the very same animal was sighted by a large group of people in the foothills of Sultanuizdag 3.0 km away from the right bank of Amu-Darya, 15 to 20 km beyond the point of first sighting in June, 1968. This suggests that tigers may still be living in several dense areas of the delta (A.M. Mambetzhumaev and A.S. Sabilaev). Оп Urta-tugai Island (about 400 km? in area, now part of Afghanistan) on the Pyandzh at Parhar, some four to five litters were discovered annually according to some accounts. This is obviously an exaggeration. 100 117 at the end of the last century and partly in the first decade of the present. Yet even toward the end of the last century their decline had begun. Changes in habitat played a decisive role only in the recent period (from the 1920’s and especially 1930’s). The main reason, however, was the extermination of the animal by military game hunters.*? Even in the 1930’s tigers bred along Pyandzh and Amu-Darya and in the lower reaches of Vakhsh and probably in other isolated places, but by 1950 (probably earlier) their population here consisted of only a few stray animals coming from the south. They lived more permanently in the lower reaches of Vakhsh, especially in Tigrovaya Balka preserve. In the 1930’s, 1940’s, and early 1950’s instances of tiger appearances were recorded in Surkhan-Darya valley at Denau and Saryassiya (1930; V.G. Heptner), the floodplains of the Kafirnigan in the north as far as Mikoyanabad (Kabadian) in the Gissar valley (killed at Rokhatinisk MTS in 1938), the Vakhsh valley in the north of Aral (Kuibyshev region), the Pyandzh, especially its upper reaches, the lower reaches of Kyzylsu in the region of Parhar, and along the river below Kolkhozabad. In the Kyzylsu tugais by 1950 there were not more than five tigers and their total number in Tadzhikistan as a whole did not exceed ten to fifteen. In the sanctuary, ‘’Пютоуауа Balka’’, tracks were seen for the last time in 1953. Thus in the region under description tigers had disappeared by the 1950’s but some stray transgressions from Afghanistan were still possible. Furthermore, even in that country the animal was very rare along the Pyandzh and Amu- Darya; and at the end of the 1960’s tigers reportedly entered Afghanistan from the Soviet Union (J. Niethammer). Between the above two sections of permanent habitation of tiger, along Amu-Darya, in a large section of the river valley the animals did not live permanently, at least not in the last 50 to 70 years, but were often sighted as intruders. Reports include tiger appearances at Kyzyl-Ayak (25 km south of Kerka), at Kerka, at Pal’vart (80 km below Kerka), and at Chardzhou. In some cases the animals may have stayed for some time and even reproduced. In the winter of 1931 a group of six tigers migrated beyond Chardzhou, down toward the Amu-Darya, and settled (? V. H.) at Dzhar- rabat-tugai (about 30 to 40 km below Darganat) and reproduced there. In the past, when the tugai and forests were plentiful and tigers abundant in the delta as well as in the upper reaches of Amu-Darya, such migrations along the river were more frequent.*> Some had in their possession several tens (!) of tiger skins (Masal’skii, 1913). *3Bogdanov, 1882; Lagofet, 1913; Masal’skii, 1913; Bil’kevich, 1924; Gladkov and Nikol’skii, 1935; Flerov, 1935; Sludskii, 1950, 1953, 1953a, and 1966; Chernyshev, 1950 and 1958; Berg and Ivanova-Berg, 1951; Pokrovskii, 1951; and others; and data of V.G. Heptner. 118 In the remote past tigers undoubtedly lived or were present on the Zeravshan, probably even in the middle of the last century, but references to this are mainly indirect (Severtsov, 1873). Along the Syr-Darya tigers were found from the estuary to the Ferghana valley. Along the right tributary of the Syr-Darya, i.e., Chirchik, they were encountered as far as Tashkent (tigers lived permanently in the immediate proximity of the town in the 1870’s) and above (reported for the Kuraminsk range 40 km south of Tashkent), and noted along the Arysu and its tributaries the Boroldai (50 km north of Chimkent) and the UI’kun-Aksu (in its upper reaches in the western spur of the Talassk Alatau). They also occurred along the entire coast of the Aral Sea where large reed jungles were abundant in the past. Tigers likewise lived in the lower reaches of the Kuvan-Darya and Dzhany-Darya. Tigers were widely distributed in Syr-Darya basin in the early and middle parts of the last century, although even then at some places, for example in the Ferghana valley, they were reported only as intruders and evidently rare. The extinction of tigers became more rapid with the appearance of Russian hunters, their population began to dwindle rapidly, and their area of distribution shrank. Tigers survived the longest in the lower reaches of Syr-Darya below Kazalinsk. Nevertheless, at the end of the nineteenth century they were evidently still met with along the river and around Chirchik, while tiger transgressions in Ferghana valley were reported even in 1903. By 1910 to 1915 below Kyzyl-Orda tigers were still quite common, but north of it encountered rarely and irregularly. Commencing from the first and especially the second decade of the present century, tigers disappeared rapidly even in the lower reaches of Syr-Darya; the last tiger there was killed in 1933 (Kazalinsk region). Some stray animals were observed even up to 1937. After an interval of several years, tigers appeared for the last time in the lower reaches of the Syr- Darya at the end of 1945, when they were observed in a section from the estuary to Solotyub (slightly above Kyzyl-Orda). These animals had come from the Amu-Darya together with wild boar, which in that year migrated in very large numbers from the Amu-Darya delta along the shore of Aral Sea to the Syr-Darya. In the middle of the winter the animals disappeared from the Syr-Darya. The arrival of tigers from the Amu-Darya was again recorded in 1924. Tigers intruded far north on the Turgai River from the estuary of the Syr-Darya and the shore of the Aral Sea. Farther east tigers were encountered along Lake Telikul’sk and in the lower reaches of the Sarysu, and lived along the Chu from the mountains to its lower reaches; the animals lived there permanently and were not uncommon even early in the present century. The last two tigers on the Chu River were killed in 1912 but resident tigers were detected even in 1929 101 119 (two pairs), 1936, and 1937. By 1940 they had disappeared (died). The tigers of the Chu River and the lower reaches of the Sarysu were linked with those of the Syr-Darya. At the beginning of the second half of the last century tigers were met with in the lower Talas and in the Biilikul’ lake system, in the northern foothills of the Karatau and Kirgiz ranges, and also evidently on the slopes of Karatau.>4 In the east tigers lived all along the Ш River up to the State boundary and throughout the system of creeks and channels in its lower reaches and delta, along the Karatal, Aksu, and Lepse, all along the reed jungles on the southern shore of Balkhash, along the Ayaguz (lower reaches), and in the rushes at Alakul. Tigers were mostly seen on the Ш, especially in the lower reaches; they lived along other rivers mainly in their lower reaches and estuaries. Even at the beginning of the present century tigers appeared only sporadically in the lower reaches of the Karatal but litters were seen even in 1929 and the last encounter (catch) occurred in 1931. On the Aksu tigers were noted in 1909, on the Lepse still in 1930, and on the Ayaguz a litter was detected in 1908. On the Ш River tigers were first exterminated in the middle course (Iliisk). In the 1890’s they were quite common there but appeared sporadically in the present century, although transgressions were reported even in 1929 and 1930. Tigers survived the longest in the lower reaches of Ili, living there until the 1930’s. There were 10 to 12 of them in 1935 and 5 or 6 in 1939. Tigers had entirely disappeared (died) on the Ш River by 1948. Evidently, even transgressions from Kuldja are hardly possible now. In the last century, even in the 1890’s tigers were rarely sighted in the hills south of the region under description. The animals were reported in the Trans-Ili Alatau, the Chu valley, the Kirgiz (Aleksandrov) range, and in particular on the southern slope to the Talasu in the Talas city region and in the Talassk valley, at Issyk-Kul’, in the upper reaches of the Naryn and on the Narynkol.* The eastern edge of the Middle Asian region of tiger habitation is formed by the Tarbagatai [range], Zaisannor [Lake] and the lower reaches of the Chernyi Irtysh, and the Kurchum valley. Here animals were encountered only in the last century, becoming rare even toward the end of it.*° 4According to data of Rychkov, 1762; М. Severtsov, 1873; Hern, 1891; Zarudnyi, 1915; Sokolov, 1924; G.V. Nikol’skii, 1930; Ognev, 1935; Sludskii, 1939, 1950, 1953, 1953a, and 1966; Kuznetsov, 1948; and others. *According to М. Severtsov (1873) the occurrence of tigers there was common and permanent. In his view they were encountered only in summer in the alpine zone. According to data of Brandt, 1856; P.P. Semenov, 1865; N. Severtsov, 1873; Finsch and Brehm, 1880*; A. Nikol’skii, 1887; Alferaki, 1891; Slovtsov, 1897; Shostak, 1927; Shnitnikov, 1936; D. Dement’ev, 1938; Sludskii, 1939, 1953, 1953a, and 1966; Kuznetsov, 1948; P.I. Semenov-Tyan-Shanskii, 1948; and others. | *Not in Literature Cited — Sci. Ed. 120 Tigers are capable of negotiating very long distances and have been seen as intruders far from the region of their normal habitation. Often they find themselves in places unfavorable for their survival, on terri- tories with an environment altogether alien to them (open steppes and deserts). In Middle Asia and Kazakhstan many distant intrusions of tigers are known. Thus, along the northern edge of the Kyzylkum and the eastern bank of the Aral Sea they crossed time and again into the lower Syr-Darya (evidently also vice versa), which is about 500 km as the crow flies. In 1945 it was conclusively established that tigers coming from the Amu- Darya to the Syr-Darya following wild boar traveled about 1,000 km in about two to three months. In the west transgressions are known in Turkmenia in the Great Balkhan (1936) and even the northern shore of the Kara-Bogaz-Gol bay (Shev- chenkovsk region, Guryev district; tigress and two cubs in 1947). In the Great Balkhan tigers undoubtedly traveled from Iran, to Kara-Bogaz-Gol, evidently along the southern Ustyurt chink [arroyo] from the Amu-Darya delta. The animals in both instances would have covered great expanses of desert. Transgressions were known from the Amu-Darya delta along the southern Ustyurt chink, and in the Ustyurt in the region of the road in Kungrad. Tigers have time and again occurred even on the southeast coast of the Aral Sea in the northern Kyzylkum and at Kuvan-Darya. From the Amu-Darya they evidently extended even to the Ural [river] estuary.°’ In Kazakhstan and adjacent parts of western Siberia tigers intruded at several places evidently from Pri-Balkhash, probably from the Chu and Sarysu Rivers, and from the Zaisan region. In the southern Altai and Pri- Altai regions tigers were reported at Altai (Katon-Karagai) station, Bukhtarminsk, Ust-Kamenogorsk, Semipalatinsk, Zmeinogorsk, Loktevsk Zavod on the Alee River, at Biisk (52°30') and even Barnaul (53°), and at other places in this same region (Setovka, В. Biisk u.; Pokrovskii Zavod). Along the Irtysh the animals reached Pavlodar and Yamyshev (about 50 km above Pavlodar). These intrusions were reported in the last century, mainly in the early part; transgressions in the southern regions were not very rare. West of Irtysh, intrusions have been recorded for the region north of Balkhash, the Karkaralinsk mountains, toward Akmolinsk, and even into the Kokchetav mountains (Sedel’nikov and Borodin, 1903; Sludskii, 1953). >7**Close to the estuary he (Yaik) was overtopped by tall reeds, where wild boar and tiger found refuge’’ (A.S. Pushkin, History of Pugachev, vol. I). This lone reference of Pushkin has no reliable confirmation in zoological literature. There is, however, a reference to the habitation of tigers in the region of nomadic Little Order Kazakhs (Brandt, 1856). These animals were sighted within the boundaries of the Ural’sk district, and in part in the Turgaisk and Syr-Daryask districts, and in Orenburg and Astrakhan provinces. 102 121 The last of these sites is about 700 km north of Balkhash. It is possible, however, that at least some of these intrusions were associated not with the habitat of the animal in the Balkhash region but with the Chu region, 1.е., Sarysu and Syr-Darya. Such distant settlements were, in the last century, facilitated to some extent by a much broader past distribution of wild boar in the north (see Volume Г of this monograph). In eastern Siberia (Fig. 50) tigers nowhere settled permanently or, if they did, it was so long ago that there exists no direct evidence. Tiger occurrences in Trans-Baikal and Cis-Baikal (see below) were, and continue to be, mere transgressions. There is no positive information about intrusions in the early part of the last century into the headwaters region of the Yenisey (Tuva, ‘‘Uryankhaisk Land’’; Radde, 1862). Fig. 50. Distribution of tigers in eastern Siberia and along the upper Amur. 1—places and years of occurrence; 2—localities in Pri-Amur (see Fig. 53); 3—references to Verkhoyansk mountain range and unconfirmed information about the upper reaches of the Irkuta, Oka, and Tuva. V.G. Heptner. — 122 Far Eastern distributional region. This area lies in the Pri-Amur and Ussuri regions; the Trans-Baikal region is zoogeographically identical to this one (Figs. 51, 52 and 53). All these regions of the range are linked with northeast China (Manchuria). The northern boundary of the habitat of the animal in this area in the historic period can only be tentatively established. Adequate information does not exist (available only from the commencement of Russian investigations, i.e., in the second half of the last century). Furthermore, multiannual fluctuations of the boundary inevitably resulted from frequent transgressions beyond the area of permanent habitation, natural causes, and disturbance by man. It is also significant that tigers lived and continue to live in the Far East not in isolated pockets along rivers in the desert as in Middle Asia, but over a large expanse with relatively uniform forest conditions. In the west, the reconstructed northern boundary of permanent habitation of the tiger commences on the Amur at the mouth of the Khumaerkhe (Kumara), flowing into Amur from the right. It runs east and southeast, encompassing the lower reaches of the Zeya and Bureya, its left tributary the Tyrma, the southern part of the Bureinsk range, the Bidzhan and Bir basins, the lower and middle reaches of the Отта (оп the north a tributary of Urma, the Berendzha) and the Kur’, and emerges at the Gorin. At the mouth of this river or slightly below it, the boundary crossed the Amur and, avoiding the Khungara basin to the south, extended toward the Tatarskii Strait. It is highly probable that the boundary actually crossed, at least at some places, slightly farther north, probably along the upper reaches of the Amguna, downstream along the Amur (at 52° in the region of Lake Kizi), and so on. South of this line tigers occurred everywhere right up to the Korean border, although absent at places in the high mountains of the Sikhote-Alin. This reconstructed range corresponds roughly to the situation obtaining in the middle of the last century. As a result mainly of direct persecution, and in part felling of forests and extermination of wild ungulates, especially wild boar, the range was greatly altered. On the whole, from the end of the last and the beginning of the present century, there has been a significant reduction of the range, mainly in the north. Thus, by 1940 the northern boundary of permanent habitation in the Ussuri region extended in general along the Iman, and the animal was almost absent throughout the northern part of the Ussuri region, along the Khor and Bikin. This reconstructed range became the region of later occurrences (but not all); as a result of reduced persecution during the war, establishment of preserves, and enforcement of measures for their conservation, the range of tiger enlarged somewhat. By 1950 tigers were found in the north along the Bikin and the southern 104 о 4 Fig. 51. Change in some range boundaries of tiger in the Far East. 1—northern boundary of permanent habitat; in part of permanent and regular intrusions in the first half of the nineteenth century; 2—approximate northern boundary in 1940, the period of utmost recession; 3— boundary in 1950; 4—some peripheral points of intrusions from 1920 to 1962; 5—from 1952 to 1962 (according to data of N.V. Rakov); 6 — points of earlier intrusions; 7— occurrences in Sakhalin (nineteenth century, no exact site). V.G. Heptner. tributaries of the Khor and along the coast reached Samarga where the animal had long been absent (Fig. 51). After 1950 enlargement of the range of tiger continued and by 1969 (data of S.P. Kucherenko and A.G. Pankrat’ev for December, 1968) comprised three sections (Fig. 52). Maximum enlargement occurred in the Sikhote-Alin. The region of permanent habitation extended there roughly Lake Orel i a | : | Е ees ЕЕ ИЕ ЕЕ a ea en ane (CIDE a a Gay’ Een ee | и — Е Е АКИ ЕЕ А Eee ee Re Vit —————— eee TY Zan! 7 Ц il | $ 138 105 Fig. 52. Distribution, nature of occurrence, and comparative numbers of tigers in Pri-Amur and Ussuri regions at the end of 1968. 1—region of permanent habitat; 2—sections of maximum population density; 3—region of regular temporary occurrence; 4— points of distant occurrence in the 1960’s; 5—direction of more or less regular migrations; 6— route of particular distant occurrence. S.P. Kucherenko and A.G. Pankrat’ev. 104 125 up to 46° М. lat. (the upper Kema); one extension ran northward on the eastern slope of the mountain range along the coast, extending slightly north of the mouth of the Samarga (about 47°30’), and another ran along the western part of the Sikhote-Alin north of the latitude of Khabarovsk (almost to 49° N. lat.). This range was surrounded by a region of temporary but regular trans- gressions. In the west this ran initially in a narrow strip along the western slopes of the Sikhote-Alin and along the Ussuri, and they continued as a broad projection to Komsomol’sk-on-Amur. In the middle of the Sikhote- Alin this region extended approximately to 48°20’ М. lat. and along the eastern slopes of the range, to 48°45’, 1.е., almost to Sovetsk Gavan’. Northward from there some individual irregular intrusions extended in the 1960’s up to the upper Tumuin, Amur at 51° N. lat., the sources and midcourse of the Gorin (Goryun; left tributary of the Amur, entering below Komsomol’sk), and the Duka River the source of the Amguna in general slightly north of 51° N. lat. The region of maximum density of the tiger population occupied almost the whole of the southern part of the region of permanent habitation, roughly from the upper Armu and upper Kolumba at the source of the Iman (slightly south of 46° N. lat.). A very small separate section of tiger habitat extended in the form of a narrow Strip, along the southernmost section of the state boundary from the level of Lake Khanka to the latitude of Vladivostok or slightly farther south. There are two regions of permanent habitation here and two of regular intrusions from the west. This section was isolated at the end of the 1960’s from the main Sikhote-Alin range described above. The third section of tiger habitat lies inside the large southern bend of the Amur. Commencing at the Amur, southwest of the town of Obluch’e, it extends initially east almost to the town of Birobidzhan and later southeast, running again toward the Amur west of the mouth of Bidzhan. The range thus encompasses the southeastern part of Bureinsk range and Pri-Amur lowland to the east. The central part of the section adjoining the Amur represents the region of permanent habitation; it is surrounded by the region of regular intrusions. Some more distant intrusions from this part of the range extended in the 1960’s to the central parts of Urma basin and the midcourse of the Kur’.°$ As in the earlier period, in the 1960’s also there were regular migrations of tiger between northeast China (Manchuria) and the Soviet Union. At the very end of this decade such migrations took place in the Amur bend, the °*By 1970 the situation in this part of the range and on the right bank of the Amur had undergone a change. This region, which invariably received migrant animals coming from beyond the Amur, is evidently devoid of tiger or is becoming so. In any case, in the winter of 1968 to 1969 not a single track was detected in Birobidzhan (A.G. Pankrat’ev). 105 106 107 126 region of the mouth of the Bikin and Ussuri, and in the section of the range in the extreme southeastern Ussuri region adjoining [lake] Khanka (S.P. Kucherenko and А.С. Pankrat’ev). The enlargement of the range described above from the time of its utmost reduction in 1940 as well as the increase in tiger population, which took place in spite of a rapid increase in human settlements and intense cultivation of the country (mainly extensive exploitation of forests), are attributed primarily to some of the conservation measures implemented after the war. In spite of improved natural conditions, which for a long time were not present in the west, tigers even here would have faced rapid extinction had such measures not been implemented. By the end of the 1960’s a fairly extensive territory was occupied by settled breeding populations and a fairly large number of regular intrusions. They are such that through very careful regulations it is possible to conserve the animals. Tiger intrusions into the Far East always were very extensive, reaching far beyond the limits of the reconstructed boundaries described above, especially in the past (Figs. 50 and 53). However, in our century and even in the past decade, tigers penetrated quite far, especially when one recalls that the northern boundary of the region of permanent habitation within the Soviet Union had receded southward markedly (see above). These tigers have been sighted in different years somewhat above the mouth of the Kumara on the Amur, at Selemdzha around 52°, north of the Berendzha River, а tributary of Urma (about 49°30’ to 50°00’; probably this is not an intrusion but a rare, more or less permanent habitation); at Niman on the right tributary of the upper Bureya (around 51°30’), on the upper Amguna (slightly south of 51°), on the Amur below the mouth of the Gorin, at Lakes Kadi and Kizi on the lower Amur, and in certain other places, extending sometimes to 54°, i.e., up to the latitude of the Amur estuary. From 1920 to 1962 tigers transgressed into an extensive region north of the Amur, as may be tentatively judged by the following reports: the region of Yerofey Pavlovich and Solov’evsk; slightly west of the town of Zeya on the Zeya; between the upper Zeya and its right tributary the Mul’shuga; the left bank of the upper Zeya at the meridian of the town of Bomnak; slightly east (on 130° E. long.) on the same latitude; the headwaters of the Nora, a right tributary of the Selemdzha; the headwaters of the Selemdzha; the right bank of the midcourse of the Amguna on the meridian of Lake Chukchagir (left bank of the Amguna); left and right banks of the Amur slightly below the mouth of the Gorin; the upper Tumnin (data of N.V. Rakov; Fig. 51). Intrusions in the 1960’s (to 1968 inclusively) for the same region were: midcourses of the Urma and Kur’, the Duka River in the headwaters of the Amguna, headwaters and midcourse of the Gorin, right bank of the Amur 106 =} р i Fig. 53. Distribution of the Amur subspecies of tiger, Panthera tigris altaica Temm. 1— range boundaries at end of the nineteenth century; 2—range boundaries by the 1930’s; 3—sections outside the range; 4—separate long dispersals of tigers. K.G. and V.K. Abramov. below Komsomol’sk, and headwaters of the Tumnin (S.P. Kucherenko and A.G. Pankrat’ev; Fig. 52). Some intrusions, partly in the remote past and partly in our day, extended significantly farther. In 1944 tiger intrusions were reported on the upper Uchur (right tributary of the Aldan) in southeastern Yakutia, approximately 57°20’ (D. Ivanov), ап4 on the Amga River (right tributary of the Aldan) at 60° N. Lat. Evidently the report of the sighting of a snow leopard on the Olekma 300 km from its mouth (Pallas, 1811; see chapter on snow leopard) also refers to a tiger. An exceptionally distant intrusion was reported 108 128 in 1905 when two tigers were encountered оп the Aldan 60 and 80 km below Ust’-Maya (about 60°45’). Less reliable is information about the appearance of tiger in the Verkhoyansk range, evidently south of it. Tiger transgressions are also known for Sakhalin.°? References to normal occurrence on Sakhalin are erroneous, certainly (Fig. 50). Evidently, the majority of these intrusions were linked with northeastern China (Manchuria) and only a portion of the eastern points were reached from the southern bend of the Amur and the Ussuri region. The distribution of tigers in Trans-Baikaliya is quite distinctive. They occur here only as intruders, evidently from the Great Khingan [mountains] in northeast China, and although they sometimes stay for quite some time, they never breed. Intrusions extend into the easternmost (often) and southern parts of the Trans-Baikaliya.°? Reports are on record of tiger at the confluence of the Shilka and Argun, at other places along the Argun, 1.е., from its mouth upstream to the Tsurukhaitui (about 50°15’), at the large Nerchinsk factory, on the Gazimur River near its confluence with the Argun (Kuchugai), at Gorbitsa on the lower course of the Shilka (about 53°05’), at Aksha on the upper Onon (near the State boundary —about 50°15’), and on the Ingoda near Uleta (about 100 km straight up the river from Chita). Intrusions into Trans-Baikaliya were not rare. Much of the information pertains, however, to the first half and middle of the last century. At the end of the nineteenth and the first half of the twentieth century, definite information about the appearance of tigers in Trans-Baikaliya is not available but without doubt intrusions did occur. However, in December, 1953 a tiger was killed at Kaktolga on the Gamimur (52°45'’—slightly above Kuchugai), another on the bank of the Shilka in the Mogochinsk region, and a third on December 29, 1953 at Balei (south of Nerchinsk on the Unda River). In that year alone eight tigers were sighted in southeastern and eastern Trans-Baikaliya. Trans-Baikaliya is evidently a region through which tiger used to penetrate even farther east- and northeastward. Thus, in 1828 a tiger was noted at Balagansk on the Angara north of Irkutsk, and tigers occurred twice in the thirties at Nizhne-Angarsk at the northern extremity of Baikal (56°30'; Sludskii, 1953a). It is possible that some of the deep intrusions mentioned above also occurred through Trans-Baikaliya (Fig. 50). °°The Far East part of the range is from Przhevalskii, 1870; Baikov, 1925 and 1927; Afanas’ev, 1934; Salmin, 1940; Kaplanov, 1948; Sludskii, 1953 and 1966; V.K. Abramov, 1962 and other works; and data of K.G. Abramov, G.F. Bromlei, S.P. Kucherenko, A.G. Pankrat’ev, and V.G. Heptner. For other tiger intrusions see the section **Biology”’. It is possible that on several occasions animals may have entered Trans-Baikaliya along the Amur valley but in general this is rarely true; concerning tiger habitat on the Amur in the expanse from the mouth of the Kumara upstream to the confluence of the Shilka with the Argun, there are no data. 109 Geographic Range outside ше Soviet Union The range outside the Soviet Union covers northern Iran, Afghanistan, much of India except the northwestern desert regions of Rajputana, Punjab, Kutch, and Sind; Sri Lanka, Indochina, Malacca, Sumatra, Java, and Bali; the Ш basin in Dzhungaria, Lobnor, Tarim and Yarkand-Darya in eastern Turkestan; northeast China in the west including the Great Khingan, and the eastern part of China. Tigers are now extinct over large areas (China, except the northeast and extreme south, much of India, Bali, etc.). If one considers range as a whole, then the tiger is, apparently, extinct over most of it.°! From the zoogeographic point of view the tiger represents a typical south Asian form, most probably with southeast Asia as its place of origin. Its present-day (historic) range is the result of colonization by two distinct groups of animals; one group via the northeast through eastern Asia, and the other via the northwest through the Near East (Iran and Caucasus) and Middle Asia. Both these directions circumvented the Central Asian highlands and Tibet. Extension into Semirech’e, Altai, and Kashgaria on the one hand, and into Trans-Baikaliya, on the other, was a secondary phenomenon; these represent the terminal areas of two groups of animals coming from opposite directions. | According to some authors the tiger as a species is associated in origin and dispersal with eastern Trans-Baikaliya, northeastern China (Manchuria), and Pri-Amur (Baikov, 1925; К.С. Abramov, 1965). The present-day range is considered the result of colonization through eastern Asia southward and through southern Siberia westward to Turkestan and the Caucasus (Mazak, 1965). This view is unacceptable not only from the zoogeographic viewpoint but also because it is based on an incorrect interpretation of the fossil remains of large Pleistocene cats of Siberia. They were long supposed tigers while, in fact, they were cave lions (Panthera spelea Gold.), which were widely distributed not only in the Palearctic but also in the Nearctic regions (N.K. Vereshchagin). Tiger remains are known only from the Upper Pleis- tocene and Holocene strata of the southern Ussuri region (Vereshchagin and Ovodov, 1968). In the Caucasus the tiger was evidently a more recent colonizer, appearing in the Holocene period (Vereshchagin, 1959). Fossil remains have not been recovered there; nor, in particular, have they been discovered in the Binagedinsk excavations. (V.G.) Geographic Variation Geographic variation of the tiger is not very sharp but nevertheless manifested. ©The reported occurrence of tiger in Sudan, with even a description of a special form, Р. с. sudanesis (Daraniyagala, 1952), lacks credibility and warrants no scientific discussion. 130 Я 7 ~~. 6 иния \) GEE SES ТТ Е ШО Я и } С | i АНИ ({ i si! , 5: ull | {| [4 0 1000 2000 3000 4000km 107 Fig. 54. Reconstructed species range of the tiger, Panthera tigris L. Territory of permanent habitation and regular, relatively short-distance intrusions. Dots indicate deep intrusions beyond the northern boundary of the range; crosses denote regions occupied or visited by tigers in the Middle Ages. V.G. Heptner. Until recently about nine subspecies were recognized, with six in continental Asia (Ellerman and Morrison-Scott, 1966). Usually three, sometimes even four subspecies are assigned to the Soviet Union. The number of names is around 20. Soviet zoologists have always recognized only a single form for the Caucasian-Turkestan part of the range. The separation of the Balkhash tiger into a distinct subspecies (trabata) has not been recognized at all. For the Far East two subspecies have long been recognized —the northern ‘‘Amur’’ (/ongipilis) and the more southern ‘‘Korean’’ (coreensis; Ognev, 1935) even though in the 1920’s it was demonstrated that these ‘‘subspecies’’ SH are merely individual variations and that only one form of tiger inhabits the Far East (Baikov, 1925). This was convincingly proved later (Stroganov, 1962). Yet even now some researchers persist in recognizing the earlier two forms of Far East tigers (Bobrinskii, Kuznetsov and Kuzyakin, 1965). The total number of tiger subspecies is evidently eight, of which two are known in the Soviet Union. By and large, the geographic variation of tiger is quite regular. The largest form comes from the most northern parts of the range of the species (Amur), while the smallest inhabits the Malay Archipelago; tigers of Turkestan, Iran, India, southern China, and Indochina occupy an intermediate position in size and some other features (type and color of coat, and extent of difference between winter and summer coats). Bergmann’s rule and insular characteristics are well exhibited. The following forms inhabit the Soviet Union. 1. Amur tiger, Р. г. altaica Temminck, 1844 (syn. amurensis, coreensis, mikadoi, mandshurica, and longipilis). Size large, exceeding on the average the size of the largest of Indian tigers. Build heavy and massive; anterior part of trunk and forelegs parti- cularly well developed. Head relatively heavy and large.*° The main background color of the winter coat is a relatively light, ocherous-yellow, with a fairly intense admixture of reddish-rust. In general, however, color is very variable and in extreme cases may be ocherous- yellow or fairly bright reddish-rust, especially on the back. In the summer coat this color is significantly brighter and more reddish. In general, the Amur tiger is one of the most variable forms of the species insofar as color is concerned.© Underside of body and inner surface of legs white and white color on flanks usually extends quite far up. White fields around eyes quite large. Dark bands vary rather significantly in number, size, form, and partly in color. Usually, they are relatively narrow, long, often bifurcate at the ends, or occur in pairs throughout. Much shorter lenticular bands, pointed at the ends, with a gap in the middle and not reaching either the sacrum or the abdomen, are frequently seen. Number of stripes relatively small, i.e., they are not close-set (Figs. 55, 56 and 57). Stripes on body black but on thighs, base of tail, and sometimes even “These features of general appearance do not lend themselves to an accurate description but are given by many taxonomists as characteristic of Siberian tiger. The long coat is partly responsible for the general impression. °“For a long time local Russian hunters distinguished two ‘‘зибзрес!е$”” of tiger— bright rust-colored and dull-colored. These two types of coloration provided the basis for differentiating the two subspecies— **Korean’’ (coreensis) and ‘*Amur”’ (/ongipilis). It was thought that bright-colored animals were smaller and their yellow underfur shorter. $ 110 Fig. 55. Color variation in the Amur tiger, Panthera tigris altaica Тетт. Typical pattern of broad, relatively sparse stripes. A—No. S 55281, Khabarovsk region, 1952/1953; B—No. S 55280, same; C— No. S 55279, same; D—No. S 29654, Amur. All specimens from the Zoological Museum, Moscow University. Photographs by D.M. Vyazhlinskii. ee тие ииеитиюр перета тете Fig. 56. Color variation in the Amur tiger, Panthera tigris altaica Temm. Rarer specimens with a pattern of relatively close-set and narrow stripes. A—No. 3029, southern Ussuri region, Sidemi River, February 20, 1884; B— No. 2980, central Amur; C—Amur-Ussuri region; D—S 50146, left tributary of central Amur (around 48°30’ М. lat.), 1940. First two specimens from collection of the Zoological Institute, Academy of Sciences, USSR and other two from the Zoological Museum, Moscow University. Photographs by D.M. Vyazhlinskii and L.G. Turova. Fig. 57. Color variation in the Amur tiger, Panthera tigris altaica Temm. More or less intermediate pattern between types shown in Figs. 55 and 56; relatively sparse bands in anterior part of body and narrow and close bands in posterior part. Left—No. S 55284, Khabarovsk region, 1952/1953; Right—No. S 29664, female, Khar River, tributary of the Pikhtsy, tributary of the Amur (49° М. lat., about 76°30’ E. long.), March, 1938. Collection of the Zoological Museum, Moscow University (see Fig. 45). flanks usually not pure black but different shades of brown or brownish. Head, middle of back, abdomen, and distal half of tail generally pure black. In animals with very vivid color, the pattern very sharp; in light-colored animals usually less sharp, especially on flanks. Moreover, pattern less distinct im animals in long-haired winter coat. Rings on tail (not counting black tip) usually eight or nine; they are generally paired (with a light-colored transverse gap along the base). In the basal region toward the top usually (more so evidently in males) a pattern in the form of angles with the apex turned forward occurs (Fig. 58). The color of subadult tigers does not differ from that of adults; tigresses at the age of several months (three or four) are paler. Seasonal coat dimorphism is extremely sharp, and a very characteristic feature of the subspecies. Summer coat hair in length and density barely differs from that of southern tigers (India), but the long and dense, even 113 113 135 Fig. 58. Predominant color type of tails in Turanian tiger, Panthera tigris virgata Matschie (left) and Amur tiger, Р. t. altaica Temm., after К.К. Flerov and I.M. Gromov, 1934. Sketch by N.N. Kondakov. slightly shaggy winter coat imparts to the Amur tiger an altogether special appearance. In summer, males, though not all, possess slightly longer hair on the neck and throat, and whiskers are faintly perceptible. In winter, however, in the majority, mainly males, the mane is greatly elongated as is the hair throughout the neck; whiskers are also very long. The tail becomes very thick and its diameter may reach 10.0 to 12.5 cm (at the base). In the summer coat the hair on the back is 15 to 17 mm long, along the top of the neck 30 to 50 mm, on the abdomen 25 to 35 mm, and on the tail 14 to 16 mm; length of whiskers 70 to 80 mm. In winter the hair on the back is 40 to 50 mm long, along the top of the neck 70 to 110 mm, on the throat 70 to 95 mm, on the chest 60 to 100 mm, on the abdomen 65 to 105 mm, and on the tail 35 to 50 mm; length of whiskers 90 to 115 mm (Mazak, 1967). The size of Amur tigers varies markedly. The range of variation among males is particularly wide; they are considerably larger in size and heavier in weight than females. It is also significant that tigers live long and continue to grow for a very long period, practically throughout their lives. Under favorable conditions the animals appear well fed and even obese. Maximum weight of the Amur tiger is 390 kg (Baikov, 1927) and 384 kg (male; V.P. Sysoev, 1952).° It is possible that these figures are somewhat exaggerated; 4A weight of 400 kg has even been reported in an exceptional case (Baikov, 1927). 136 in any case, they pertain to large old animals, which are rare at present. The existence of animals weighing 325, 340, 350, and 360 kg has, however, been established (Baikov, 1925; Bel’skii and Bromlei, 1953; Mazak, 1965, cited from Sludskii, 1966). Males weighing up to 320 kg and females up to 180 kg are considered a fairly ‘“‘normal’’ maximum (Baikov, 1925). This figure is probably high for females, however. Some individual animals, as pointed out, greatly exceed this maximum. This is particularly true of the past, when tigers were bigger and lived longer. Early in the present century, ‘а splendid male of moderate size, well fed, and even obese . . . probably attained 160* kg’’ (its total length was about 360 cm—‘‘18 chetvert’’; Baikov, 1927). The maximum known body length of Siberian tiger is 317 cm (along the curve of the back). With a tail length of about 100 cm, this giant measured a total length of slightly less than 420 cm (Barclay, 1915; cited from Mazak and Volf, 1967). A body length of 300 cm and a total length of 375 cm (1.е., body length about 275 cm [sic]; a tiger of ‘colossal measurements”’; Baikov, 1927), have also been reported. Based on these data the ‘‘normal”’ maximum size of Amur tiger is as follows: body length (along curve) of males up to 290 cm and of females to 190 to 200 cm (probably slightly more); tail length to 115 cm; length of hind foot in males to 40 cm or slightly more and in females to 35 cm or somewhat more; transverse length of ears in males to 120 mm or more and in females about 100 mm; and height of males at shoulders to 115 or 116 cm, possibly even more.® Data on several animals as measured by zoologists ‘Чт the flesh’’ are given in Table 1. * Apparent error in original Russian; should read 460?—Sci. Ed. Height at the shoulders is generally not stated by authors. An old male with the legs stretched (as suggested for measurements) reached 124 cm (Mazak, 1967). For the big tiger referred to above with a body length of 317 cm, the height at the shoulders was given as 140 cm (Mazak and Volf, 1967), which hardly seems likely. A critical analysis of some animals and data for specimens collected have been given by S.U. Stroganov (1962). The figures given in that work, published posthumously, are contradictory and partly erroneous (pp. 428 and 437, and others). The following are some additional published data on sizes of Amur tiger: weight of adult males 196 and 217 kg; male three-year-old, 115 kg (С.Е. Bromlei); female three to four years old from central Amur (in Radde region), body length 178 cm and tail length 87 cm; male caught slightly downstream along the Amur, 180-185 cm and 93-94 cm (Radde, 1862); male of ‘‘colossal’’ dimensions (killed in 1911), length with tail 375 cm, height at shoulders 116 cm, and weight of ‘‘cleaned’’ animal, 250 kg (Baikov, 1927); animal caught in the winter of 1927/1928 in Kirin province (northeast China), body length with tail 400 cm and weight 325 kg (Baikov, 1929); female killed on December 16, 1927 near Modaoshi station on the Chinese-Eastern railway, body length 295 cm and weight 200 kg (usual weight of tigers 160 to 200 kg; Baikov, 1928); male killed in 1953 in Trans-Baikal, body length 200 cm and weight 195 kg; another from there 250 kg (Korneev, 1954); two tigers killed in Primor’e, body length with tail 325 and 310 cm; animal from around Ussuri, body length 284 cm, height at withers (Contd.) Г 116 117 137 The skull of the Amur tiger is distinguished not only in its large overall size, but also in the great development of crests, especially the sagittal; the height and strength of the latter exceed those of other forms of tiger and the lion. Skull measurements are given in Tables 2 and 3.° Found in Ussuri and Amur regions and Trans-Baikaliya, and as a vagrant in Pri-Baikal and southern Yakutiya. Outside the Soviet Union, in the northern and eastern parts of northeast China (Manchuria), the Korean peninsula, and as a vagrant in the northeastern extremity of the Mongolian People’s Republic. 2. Turanian tiger, Р. Е virgata Nliger [sic; Illiger], 1815 (syn. septentrionalis, lecaqui [51с; lecoqui}, and trabata). Compared with the Amur tiger, general build somewhat less massive and size, on the average, slightly less. Main background color of the skin variable but in general a brighter and more uniform reddish-rust, especially the summer coat, but evidently bright even in winter (Fig. 59). Dark stripes narrower, long, fuller, and closer set. Color of stripes a fairly distinct admixture of brown or cinnamon shades, sometimes even an ‘‘intense cinnamon’’ (Ognev, 1935).°’ Pattern invariably pure black only on head, neck, middle of back, and at end of tail. Rings on tail usually arranged singly at least in terminal half; angular patterns at base of tail less developed than in the Amur tiger (GES II 59). Seasonal color dimorphism prominent but less sharp than in Amur tiger. Winter coat considerably lighter in color and paler than summer coat with a less distinct pattern. Hair on winter coat much longer and denser and coat may even be ‘‘shaggy’’. Length of hair on back in summer 8 to 13 mm and on abdomen 20 to 30 mm; on nape, hair up to 20 to 50, and even 90 (footnote 65 contd.) 85 cm, and weight 221 kg (Antonov, 1950); another, body length 225 cm and weight 112 kg; maximum dimensions of *‘Amur’’ tiger, body length 276 cm and tail 100 cm; ‘*Korean”’ tiger, 300 cm and 80 cm; and weight of Amur and Korean tigers up to 340 kg (Baikov, 1929; from data furnished by A.A. Sludskii). A large male caught in its first year in the wild and raised in a zoological garden (Prague) weighed about 250 to 260 kg (Mazak, 1967). A group of three animals in a zoological garden weighed: 11-year-old male, about 317 kg; 8-year-old female, 286 kg; and 3-year-old male, about 299 kg (Mazak, 1967). These figures are erroneous, at least in the case of the female and young male, and constitute exaggerations. "The sizes of almost all the skulls preserved in Soviet museums are given in the Tables; measurements which fall slightly beyond the indicated limits are not included. 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Skull measurements of female Amur tigers (from V. Mazak, 1967) i ao aa с Old adults! Measurements min—max Greatest length 8 279.7-310.2 296.9 Condylobasal length 7 252.2-273.4 266.1 Basilar length 7 233.4-252.7 247.2 Rostral width 9 76.4-86.5 82.8 Zygomatic width 9 190-203.6 196.3 Interorbital width 9 52.3—59.2 56.1 Pestorbital width 9 56.3—60.4 58.3 Mastoid width 8 112.2-121.6 118.3 Length of lower jaw й 188—201.8 196.2 Length of upper carnassial tooth |9 31.9-34.9 32.9 Length of upper tooth row 9 85-93 у 90.7 Note: Materials and measurements the same as shown in Table 2. ‘In У. Mazak’s Table 1 (pp. 554 and 555) the animals are shown as old; in Table 3 (p. 558) as adults. The former has been used here since it would appear to be more correct, judging from skull measurements (V.H.). mm long. In winter, hair on back may reach 30 mm or more in length. In this subspecies hair evidently tends to grow much longer along the top of the neck (mane) and on the cheeks (whiskers). Data on the size and weight of Turkestan tiger are scanty. Maximum body length of males exceeds 200 cm, reaching 213, 224, and 217 cm, with a corresponding total length of 312, 324, and 309 cm, perhaps even more (see below; a body length of 270 cm has been recorded but probably was 284 cm measured as a skin). Females smaller in size; however apparently their body length may reach 200 cm (total length, 267 cm) but normally ranges from 160 to 180 cm (total length 240 to 260 cm). The maximum known weight exceeds 240 kg but evidently could be greater. Thus, although the Turkestan tiger is generally smaller than the Amur, it nonetheless attains large proportions. The following information is available on the dimensions of some °8Conceming Trans-Caucasian tigers, К.А. Satunin (1915) reports: ‘АП winter skins examined by me had very long and dense fur of dirty-yellow-light-brown color with indistinct dark stripes.’’ “‘In winter (the fur) is very long and shaggy’’. ‘‘The very longest fur . . . was that of a huge tiger killed at the end of February 1899 in Prishibinsk section (Lenkoran; V.H.). Its fur was very dense and long but coarse and had some indefinite dirty-yellow-gray or brown color with a barely perceptible dark pattern; a reddish shade was not discernible at all . . . [I saw no] other animal with such long fur of a dull color’’. ‘°... The shorter (the fur), the more vivid the reddish shade and the sharper the black stripes.”’ 116 Fig. 59. Variation in color of Turanian tiger, Panthera tigris virgata Matschie. 1 — №. 3027, **Kirgiz steppe’’ (probably Semirech’e— V.H.); 2—No. $ 33151, lower Ili, about 250 km above mouth, November, 1939; 3—No. 2983, Lobnor (eastern Turkestan— V.H.), November, 1889; 4— No. 8679, Ferghana; 5—No. 2981, *“Turkestan,”’ 1872; 6 — №. $ 52871; upper Pyandzh, Tadzhikistan, March 30, 1950. Nos. 1, 3, 4, and 5— pelts from collection of the Zoological Institute, Academy of Sciences, USSR; Nos. 2 and 6—from collection of the Zoological Museum, Moscow University. Photographs by D.M. Vyazhlinskii and L.G. Turova. 118 143 individual animals. Body length 162 cm, tail length 86.5 cm, total length 248.5 cm (Altai, on Aleya River; Spasskii, 1820); male—body length 181 cm, tail length 98 cm, total length 269 cm, weight 181 kg (southern Balkhash, 1930; A.A. Sludskii); female—body length 250 cm, tail length 67.5 cm, total length 267.5 cm (Kirgiz range; Chekal’shchik); body length 172 cm, tail length 91 cm, total length 263 cm, weight 132 kg (Biisk district, 1839); male—length from ears (!) to base of tail 194 cm and tail length 89 cm (actual body length was evidently about 220 cm, total length about 309 cm; V.H.) (Syr-Darya around Karmakchaya, G-ii, 1887); body length more than 284 cm”, weight more than 240 kg (lower Ili, 1891, T.V., 1910); total length about 306 to 340 cm (‘‘17 chetvert’’; Kerki in the upper Amu-Darya; Gavrilov, 1896); total length 270 to 300 cm (‘‘15 chetvert’’; mouth of Amu- Darya; Kosarev, 1899) body length 217 cm (Kungrad, Amu-Darya delta; Kuznetsov, 1896)’!; body length 224 cm, tail length 100 cm, total length 324 cm (Ferghana, 1903; Berg and Ivanova-Berg, 1951); female—total length 259 cm (Pocock, 1929; cited by Mazak, 1965); adult male killed on January 10, 1954 at Kaine-Kasyr on the Sumbar (southwest Kopet-Dag)—body length 225 cm, tail length 105 cm (Shukurov, 1958); adult female—body length 165 cm, tail length 95 cm, total length 260 cm, weight 135 kg (upper Pyandzh, Tadzhikistan; March 30, 1950); adult female—body length 165 cm, tail length 76.5 cm, total length 241.5 cm, length of ears 11.0 cm, length of hind foot 31.5 cm, weight 97 kg (Kyzylsu River, Tadzhikistan; June 26, 1950; Chernyshev, 1958); small tigress (‘“Theresa’’) acquired for Moscow Zoological Garden in 1924, a subadult from Iran which died in 1942—body length 160 cm, tail length 86 cm, total length 246 cm, length of ears 11 cm, length of hind foot 32 cm, weight 85 kg (Zoological Museum, Moscow University); male—body length 187 cm, tail length 90 cm, total length 277 ст (Lenkoran’; skin measurements); female—body length 178 cm (Lenkoran’; skin); female—body length 157.5 cm, height at shoulders 78 cm (Lenkoran’; Dinnik, 1914); body length 173 cm, tail length 110 cm, total length 283 cm (Trans-Caucasus: mounted skin); body length 161 cm, tail length 93 cm, total length 254 cm (Trans-Caucasus; mounted skin); male—body length 270 cm, tail length 90 cm, total length 360 cm (‘Визе tiger’’ from around Prishib, Lenkoran’; February, 1899; skin just removed), Satunin, 1915.72 Maximum skull length in males 297.0 to 365.8 (385.0—V.H.) and female— 195.7 to 255.5 mm; condylobasal length in males 259.0 to 307.9 Material gathered and furnished by A.A. Sludskii. Probably measured from skin or total length (V.H.). After Ivanova-Berg (1927). This account about exhausts the information available on measurements of tiger of this subspecies, nowadays almost extinct. 144 and females 225.0 to 263.2 mm; zygomatic width in males 219.0 to 254.0 and females 183.0 to 203.2 mm (Mazak, 1965). As mentioned above, the dimensions of the Turkestan tiger are somewhat less than those of the Amur tiger, but this is true more with respect to mean dimensions of the general population. Large animals conforming to the above-mentioned maximum proportions of Amur tiger are not known in the population of the Turkestan form and are evidently absent. However, very large individuals are also found among the latter. Such for example is the Prishibinsk tiger listed above with a body length of about 270 cm.” A tiger, killed on the Sumbar in Kopet-Dag on January 10, 1954 (stuffed skin in Ashkhabad Museum) has a body length of 225 cm and a greatest skull length of 385 mm; these measurements are considerably more than the known maximum for this subspecies and even slightly exceed the maximum for the Amur subspecies (see Table 2). The condylobasal length of this skull is 305 mm and zygomatic width 205 mm. The canine along the front curvature is 70 mm long, which is the maximum for Russian tigers (V.G. Heptner). The skull of the animal from Lenkoran’ (Belyasuvar) has a maximum length of 362 mm, zygomatic width of 248 mm, and interorbital width of 78 mm (Satunin, 1905). This form is found in Middle Asia and southern Kazakhstan, in the east (as vagrants) to the western foothills of the Altai and eastern Trans-Caucasus. By the end of 1950’s it had become extinct in Soviet territory and only some stray animals occur irregularly as intruders in the south-eastern Trans- Caucasus, southern Turkmenia, and southern Tadzhikistan (Amu-Darya). Outside the Soviet Union it occurs in the northern parts of Iran and Afghanistan, western Dzhungaria, Tarim Basin, and Lobnor (Kashgariya). The Turkestan tiger is a well-defined form which differs from the Amur tiger in several features. At the same time it has more features in common with the Indian tiger (P. ¢. tigris), evidently, than with the Amur tiger. This is reflected in general dimensions and in the nature of the coat. The summer coat in density and length of hair is similar to that of Indian animals, while the winter coat is longer and denser. The general shade of the coat is also similar but the stripes of the Turkestan subspecies are usually narrower, longer, and closer set. The color of the stripes can hardly be called pure black, as cinnamon or brownish tones predominate. $: 2 + Outside the Soviet Union, the following subspecies are recognized: 1) Chinese tiger, Р. t. amoyensis Hilzh., 1905—central and southern parts of 3К.А. Satunin (1905) reports that it was ‘‘а tiger of immense proportions . . . 1 saw it in the flesh. It appeared to me to be no smaller than the common Tuzemna horse’’. 119 145 eastern China, except the extreme south; 2) Indochinese tiger, Р. t. corbetti Mazak, 1968—extreme southeast of China (Hunan, Kwangsi, Kwangtung provinces), Vietnam, lower Burma, Malacca, and evidently Cambodia, Laos, Thailand, and northeastern parts of upper Burma’*; 3) Indian tiger, P. 1. tigris Linn., 1758 — ша, Assam; limits and relationship to preceding form not known; 4) Sumatran tiger, Р. t. sumatrae Pocock, 1929—Sumatra; 5) Javan tiger, Р. t. sondaica Temm., 1845-—Java; and 6) Balinese tiger, P. t. balica Schwarz, 1912, —ВаН.” (V.H.). Biology Population. In the extreme northwestern part of the range and in Trans- Caucasus in the early eighteenth century tigers were still found in small numbers on the east coast of the Black Sea in the Kolkhid lowland and in Adzharia. They became extinct there in the first half of the nineteenth century. In the last century and early in the present one, tigers were seen from time to time in eastern Georgia—the last animal was killed in 1922, and in Armenia where one was caught in 1948 (Sludskii, 1966). In the middle of the nineteenth century in extreme southern Azerbaidzhan—in the Lenkoran’ lowland and Talysh mountains, tigers still survived in significant numbers and also reproduced there. In the middle of the last century 10 to 20 tigers were caught around Lenkoran’ every year in autumn and winter (Brandt, 1856). In 1896 tigers were ‘‘not particularly rare’’ in Lenkoran’ and Talysh regions and one or two animals were still being killed annually (Dinnik, 1914). In the present century the tiger population in Trans-Caucasus decreased rapidly and the animals seen there only periodically as intruders from Iran. Only at Talysh do tigers still live more or less permanently. In Trans- Caucasus only two tigers were caught between 1920 and 1950 and about ten between 1950 and 1966 (Sludskii, 1966), including one in 1961 near Astara and two between 1963 and 1966 in the Lenkoran’ region. In Talysh only single individuals have been encountered (mid-1960’s; Lagidze, 1964; Aliev and Nasibov, 1966). In Turkmenia in the last century tigers were fairly common in Tedzhen. Here they were encountered in the lowlands even in the 1880’s and 1890's. Along the Murgab in the past they prowled in sufficient numbers but toward the end of the century were common only in the very headwaters and along the Kushka River. Tigers did not live in Kopet-Dag, but only entered there from Iran. They were frequently seen on the Atrek, Sumbar, and Chandyr “The relationship of this form to neighboring ones, especially the Chinese form, requires further confirmation. 5 66 : : . . 3° See *‘Geographic Range outside the Soviet Union’’, р. 129. 119 120 146 Fig. 60. Habitat of tiger and wild boar in Semirech’e, old river bed of the Ili, Est-Bakanas River in the delta. Photograph by V. Marakov. in western Kopet-Dag. Along the Amu-Darya in the last century the animal was known everywhere; by the early twentieth century it was already a rare vagrant mainly in the upper reaches and extreme lowlands (V.G. Heptner). By the beginning of the present century in Turkmenia, tigers had become almost extinct except on the Amu-Darya. In the 1930’s its population somewhat increased but not for long. From 1900 through 1968 only ten tigers were caught in all; of these, the last was killed in 1954 in western Kopet-Dag at Kaine-Kasyr (Heptner, 1956; Shukurov, 1958; Sludskii, 1966). The occurrence of tigers in Turkmenia is possible, and from farther in Iran and Afghanistan. In Uzbekistan tigers were comparatively rare even at the end of the last century. In the 1930’s their number increased slightly; for example in the lower reaches of the Amu-Darya 12 to 15 animals were counted in 1942. Later, in spite of a ban on hunting, their population decreased rapidly. In 1947 in the delta of this river, there were only five or six animals and the last was killed in 1950 or 1951.7 See footnote no. 50 above, р. 116. 120 147 Fig. 61. Reeds in places formerly inhabited by tiger in Balkhash. Photograph by S.V. Marakov. In Tadzhikistan tigers were still common in the 1930’s in the Pyandzh, Vakhsh, Kyzylsu, and other valleys, but are now extinct. In the early 1950’s the number of tigers in Tadzhikistan would not have exceeded 10 to 15 animals (Chernyshev, 1958), but probably Chernyshev erred and the number of animals far less. From 1942 through 1950 in Tadzhikistan only 10 animals were caught. In this republic tigers disappeared completely in 1953 or 1955. 121 148 There is only one report, which needs confirmation, that the predator appeared in the Pyandzh valley in the Chubek region in 1963 (A.A. Sludskii). In Tadzhikistan tiger intrusions from Afghanistan are still possible. In Kirgizia, even in the last century, tigers were rare in the Issyk-Kul’ region, in the Tersk Alatau, in the Chu valley, and along the Bol’shoi Kabin, and disappeared there roughly in the 1880’s. South of this country they became extinct in the early 1890’s (Sludskii, 1966). In the last century tigers were still common in Kazakhstan, in the Syr- Darya, Chu, Ili, Karatal, and other valleys, in Balkhash, Alakul; and Zaisan from where they intruded from time to time northward to Kokchetav, Pavlodar, Barnaul, and to the Ishim and Irtysh. In the Tien Shan tigers were regularly found, but were very rare in the southern Altai. In the last century and early part of the present tigers periodically occurred in Ustyurt, having entered there from the lower Amu-Darya. They were often encountered in the valley of the Syr-Darya. “‘They (tigers ——A.S.) were found in large numbers in reeds around the Aral Sea and along the Syr-Darya and, occurring there together with humans, perpetrated serious attack on camels and horses”’ (Rychkov, 1762). Already in the 1840’s and 1850’s the tiger population in Syr-Darya valley had begun to decline quickly, due to killing of the predators themselves Fig. 62. Reed thicket 4.0 to 5.0 m high in tiger habitat in Semirech’e. Dzhel’- Turanga, in the lower Ili. November, 1959. Photograph by A.A. Sludskit. 121 4 149 and of wild boar, their main source of food, and destruction of the reeds. At that time ““hunting teams’’, some officers and Cossacks from the local garrisons, killed 8 to 12 tigers in a brief period. In the early twentieth century in Syr-Darya this predator survived only at a few places. In five years (from 1905) in the Syrdar’insk district only nine tigers were killed (Zarudnyi, 1915) and their population continued to decline. The last two tigers were killed on the Syr-Darya, in 1924 in the Aral region, and in 1933 in the Kazalinsk region. Tiger tracks were, however, noticed even in 1937 and 1945 when some animals entered into the lower reaches of the Syr-Darya from the Amu-Darya delta (Sludskii, 1966). In the Chu valley tigers were extinct by 1912 or at least 1916-1917. In the Ili delta, where tigers had survived for a particularly long period, only ten were found in 1935. The last was killed here in the winter of 1934/1935 and the animal had disappeared totally from southern Pri-Balkhash in 1948. In the Zaisan depression tigers were already extinct in 1894. Thus tigers are exceptionally rare nowadays and seen only as vagrants in the Trans-Caucasus; they have disappeared completely from all the republics of Middle Asia and Kazakhstan. Intrusions are still possible into Tadzhikistan and extreme southwestern Turkmenia. Fig. 63. Upper Ili delta, habitat of tiger and wild boar. Roe deer were also very common in this region. Dzhel’-Turanga, Semirech’e. Photograph by V.S. Marakov. 150 In eastern Siberia in the first half of the nineteenth century tigers were regularly seen at Baikal (Ehrenberg, 1831; Simashko, 1851; Sel’skii, 1856) and in southeastern Trans-Baikaliya, intruding from there far to the north, appearing from time to time even in the Verkhoyansk range (Silant’ev, 1919). Although not every year, one or two animals were caught in Trans-Baikaliya in some years, and tigers were caught from time to time even in the early twentieth century. Thus some were killed in 1912, 1926, and 1928. After a long interval they reappeared in Trans-Baikaliya, in 1953 and 1954. In 1953 some eight animals were counted, of which two were caught (Sludskii, 1966). From 1963 to 1966 tracks of the predator were found on the Uryumkan and Gazimur Rivers (Sinitsyn, 1966). Tigers penetrated into Trans-Baikaliya from the Great Khingan range of China. Farther east, in Amur basin, tigers were encountered in the last century and early part of the present one fairly regularly northward to 50° N. lat. They were common south of the Bureinskiy range on the Amur around 49° N. lat. (Radde, 1862). In the 1890’s in the Ussuri region about 120 to 150 tigers were caught annually (Silant’ev, 1898). Early in the present century the tiger population in the Far East declined rapidly, and by 1912, for example, only two animals were caught in the Amur region, and 55 in the Fig. 64. Tugais of turanga and tamarisk at Iman-baba, site of former tiger habitat in the Murgab valley. Turkmenia. April, 1962. Photograph by A.A. Sludskii. 151 Ритог’е (Baikov, 1927). The tiger population suffered a particularly great reduction in the Far East in 1929. Collectivization of agriculture and, later, measures undertaken for the conservation of tigers resulted in an increase in their population so that in the 1940’s the animal was seen in regions whence it had disappeared long ago. Conditions for the reproduction of tigers remained favorable in the Far East until 1948; subsequently intense netting of cubs and shooting of adult animals decimated the population. Only 27 cubs were netted between 1930 and 1938 in the region under description (Kaplanov, 1948), while 56 were caught in the eight-year span between 1948 and 1956. From 1950 through 1960 in the Primor’e region alone (southern half of the Ussuri) 23 tigers were killed. The Game Inspectorate studied the tiger population in the Khabarovsk and Primor’e regions in 1956 and 1957. In the Khabarovsk region the area of tiger habitat was about 80,000 km?. The occurrence of 28 tigers was established over an area of 56,000 km. In the Primor’e region 35 animals were counted in an area of 21,000 km2. In 1958 and 1959 in the Primor’e region an expedition of the Far East 122 Fig. 65. Stream and tugais in the Tigrovaya Balka preserve. Habitat of tiger, wild boar, Bukhara deer, and jungle cat. Lower Vakhsh, Tadzhikistan, October, 1960. Photograph by A.A. Sludskii. 152 Branch of the Academy of Sciences of the USSR, and the Central Game Board of the Russian Soviet Federated Socialist Republic estimated the tiger population at 55 animals. These studies did not cover certain regions in which 8 to 10 or more animals lived. Evidently, in the whole of the Primor’e region 60 to 65 tigers lived in the first half of 1959. Their range covered 23 administrative regions with a total area of 51,000 km? (about 30% of the whole area of the region). In the Khabarovsk region (data of the Board of Game and Hunting) there were some 35 tigers in 1959 and a few more confined to the Amur basin. Thus in the first half of 1959 there were about 100 tigers living in the Soviet Far East, constituting their total population for all of the USSR. The range of the tiger has undergone some changes in recent years in the Far East. As a result of a sharp reduction in population of wild boar, the predator has disappeared in the Amur region. In the Primor’e and a series of portions in the northern part of the tiger’s range—in the upper Iman and Bikin, the population has declined somewhat. This reduction is explained, on the one hand, by the further human colonization of forest massifs along the Armu and Tatibe Rivers and the Iman basin, and on the other, by a great Fig. 66. Thickets of turanga, tamarisk, and plume grass, the site of tiger habitat in Tigrovaya Balka preserve. Lower Vakhsh, Tadzhikistan. October, 1960. Photograph by A.A. Sludskii. 153 reduction in the population of wild boar in these regions due to inadequate yields from food crops and winters of intense snowfall for several consecutive years. However, in southern and southeastern regions of the Primor’e the tiger population has risen even in cultivated regions adjoining the Suchan, Maikhe, Tudagou, and other rivers; the animal is now encountered more frequently than in 1959. The total tiger population in the Soviet Far East, and thus for the Soviet Union as a whole, consisted of roughly 120 animals on January 1, 1965 (Sludskii, 1966).”7 In the present century the tiger population has decreased in countries adjoining the Soviet Union. Thus, in the regions of Turkey adjoining Georgia and Armenia several tigers were caught every year in the middle of the last century (Blit, 1863). At the present time tigers are extinct in Turkey. They have been preserved in northern Iran along the eastern slope of the Talysh and on the western banks of the Caspian [Sea], and they also inhabit Mazanderan, Gilan, Gorgan, and Khurasan provinces. In the 1930’s in Iran 80 to 100 tigers still survived but subsequently these numbers have declined. Tigers became “‘quite rare’’ in the forests on the southern coasts of the Caspian. Apparently intense felling of forests has caused the animal to disappear altogether from Iran in recent years (Misonne, 1959; Ley, 1967). In Afghanistan, before the 1950’s, tigers were common in the tugais along the left bank of the Pyandzh, from whence they frequently intruded into Tadzhikistan. In the last decade such intrusions have ceased, signifying, possibly, the disappearance of the animal from the former region. In the last century tigers were found in thickets along the Gerirud, Kunduz, and Murgab rivers, but today they have evidently disappeared there also. In China, Dzhungaria, and Kashgariya in the last century tigers were fairly common in the Tarim depression, along the Tarim itself, around Lobnor, and along the Khotan, Yarkand, Kyzylsu, Kashgar, Manas, Urungu and other rivers, and in the Tien Shan. They had disappeared from the Tarim basin even in the 1920’s and from the Tien Shan at about the same time. In 1959 tigers occurred with great rarity only along the Manas River, being absent already in other regions of Sinkiang; evidently at present they have now disappeared from there as well. Farther east, in Hailungjiang province (northeast China), adjoining the Soviet Union, and in the more southerly Kirin province, there were still 200 ™n early 1968 the total tiger population in the Far East was estimated at 122 to 148 animals. The relatively poor increase in population, apart from other factors, is explained by illegal shooting in the remote regions. In 1966 to 1967 about 20 tigers were reportedly killed illegally (Kucherenko, 1970). Nevertheless the ban on hunting and enforcement of conservation measures have tripled the pre-1940 population level of the animal in the Far East (V.H.). 154 Fig. 67. Turanga valley forest (tugai). Habitat of tiger. Lower Vakhsh. Tadzhikistan. Tigrovaya Balka preserve. April, 1966. Photograph by G.N. Sapozhnikov. to 250 tigers in 1958. As a result of intense felling of forests their population declined rapidly in this region. In the latter half of the last century about 150 tiger skins were exported annually from the Korean Peninsula. Today in the South Korea they are seen no more. In North Korea they have survived in regions adjoining Kirin province of the Chinese People’s Republic; some 40 to 50 were supposedly still living there in the 1950’s (Sludskii, 1966). At present world tiger reserves number 15,000 animals (Perry, 1964). Countrywise the distribution is as follows: Soviet Union—120; Пап— 80 to 100; possibly less; India and Pakistan—3,000 to 4,000; Chinese People’s Republic—2,000; Korean People’s Democratic Republic —40 to 50; and Malaysia—3,000. There is no information for other countries (Sludskii, 1966). Habitat. In the southeast Trans-Caucasus tigers were mostly confined to forests in Talysh lowlands in places where reed thickets along marine lagoons (mortsov) and streams were adjacent. These dense forests with abundant prickly vines could be negotiated only with the help of an axe 124 155 Fig. 68. Thicket of giant plume grass (Erianthus purpurascens) at site of tiger habitat in “‘Tigrovaya Balka’’ preserve. Lower Vakhsh and Pyandzha. October, 1960. Photograph by А.А. Sludskii. or scythe. Wild boar flourished in both forests and reeds but tigers pursuing them were encountered only rarely in the montane forests of Talysh. In Turkmenia, Uzbekistan and Tadzhikistan the main habitats of this predator are drainage basins of rivers and lakes, densely grown reeds attaining gigantic heights (5.0 to 6.0 m), and plume grass or tugai forests consisting of poplar, oleaster, and willow. All tugai trees are entwined with vines [Clematis]. In the lower stratum, cane, reed grass, licorice, camel’s needles, and other plants flourish. These places abound in wild boar today, and formerly Bukhara deer were also common. At such places, where the approach to the river is across long expanses of solonchaks or sands, tigers Fig. 69. Imbishi area in Sudzukhin preserve — а common place for tiger residence. Nut pine-broadleaved taiga rich in Manchurian wapiti and wild boar. Southern Ussuri territory. December, 1964. Photograph by У.Е. Prisyazhnyuk. are absent or encountered only occasionally. Rarely these animals are seen in the foothills and the lower montane belt along valleys and gorges overgrown with pistachio, juniper, dog rose, etc. In Tadzhikistan tigers ascended to higher elevation from below extremely rarely. In some regions tigers lived fairly regularly close to human settlements. Relatively small reed thickets, covering several hundred hectares set between auls [villages] where wild boar roamed, were suitable for tiger. For example, in the Yakhsu River valley in Tadzhikistan, long inhabited by man with considerable land under pasturage, tigers lived permanently in the thickets in sections 1.0 to 3.0 km long and about 500 m wide; sections were located between rice and cotton fields. In the lower Syr-Darya and along rivers and large lakes in southern Pri-Balkhash, tigers lived in extensive reed floodplains alternating with lakes, streams, and series of sand knolls overgrown with saxaul, dzhuzgun [Calligonum |, winterfat, and other plants, and also in the tugais. These areas abounded in wild boar and roe deer (in Syr-Darya, Bukhara deer replaced roe deer). 126 _ 129 5 Fig. 70. Montane taiga in the Tun’shi valley. Habitat of tiger. Sikhote-Alin preserve, Ussuri region. August, 1964. Photograph by У.Е. Prisyazhnyuk. In Kazakhstan, and also in Kirgizia, tigers were sometimes encountered in montane belts, in summer ascending up to the permanent snow line. In the Kirgiz, Trans-Ili, and Dzhungarsk Alatau [ranges] they were caught in fir and juniper groves at heights of 2,500 to 3,000 m above sea level. In the mountains tigers were found even along valleys in thickets of tamarisk, willow, or sea buckthorn. These predators ascended the mountains in pursuit of wild boar and domesticated cattle which migrated there in summer. In Middle Asia and Kazakhstan habitats preferred by tiger roughly comply with the following requirements: 1) abundance of wild boar and Bukhara deer which serve as their main prey; 2) abundant available water, which is important for this predator that drinks several times a day; 3) availability of dense, impenetrable thickets in which tigers set up dens; and 4) low snow cover. These ecological requirements of tiger explain their absence in deserts far from water sources and only temporary or chance appearances in steppes and at high altitudes in hills. Tigers live in the Far East under a different set of conditions. Here they are mostly confined to low mountains, especially in the last decade. In the last century they were even encountered at low altitudes but soon were displaced by man. Nowadays the most common habitats of this predator are montane river valleys and pad’s overgrown with Manchurian-type 127 Fig. 71. Ta-Chingou region and Taman [Foggy] mountain. Habitat of sika, Manchurian wapiti, wild boar, and goral; an area in which tigers regularly hunt. Here in the fog, the animal catches wapiti and sika coming out to sun themselves. Sudzukhin preserve, April, 1967. Photograph by V.E. Prisyazhnyuk. vegetation with an abundance of nut pine and oak. The animal also lives in pure nut pine forests, and also among mountains overgrown with deciduous shrubs, or in oak or nut-tree groves. Tigers only travel through dense spruce forests and do not remain in them. While tigers are attracted to river valleys and nut pine forests by wild boar, they mainly hunt abundant wapiti and moose in burned-over clearings (Kaplanov, 1948). Tigers are particularly attracted to rocky sections. ‘“Permanent habitations are invariably found at places where mountain ranges are inaccessible, slopes very steep and precipitous, places with extensive rock debris, overhanging cliffs, rocky crests, niches, and caves’’ (Baikov, 1925). Tigers can readily hide in rocky sections and, what is more important, there is less snow in rocky sections in winter. While hunting and wandering tigers prowl through quite high mountains, ascending to 1,600 m above sea level, nevertheless they prefer to move along gentle slopes and in low saddles. During times of movement the animal is seen in quite diverse biotopes, including sea coasts, hayfields, shew 159 etc. Quite often they are confined close to villages, especially in winter when it is difficult to capture food. In the Primor’e and other parts of the Far East where frequent and heavy snowfalls occur, tigers avoid sections with a deep snow cover since the availability of ungulates in such areas is poor. Moreover, an unstable crust hinders hunting. On thin crusts the tiger’s movement is audible and frightens wild animals; furthermore, the predator is apt to hurt its legs. The legs of a female tiger killed at the end of January, 1868 in the Ussuri region were severely injured. The fur “‘on the inner side of the front and hind legs were worn away and the skin covered with bleeding sores’’ (Przheval’skii, 1870). The heavy bodied tiger moves with difficulty in deep snow cover and breakable snow crust. The weight load per 1.0 cm? of resting paw surface in Primor’e is equal to 158 g; among our wild cats this predator exerts the highest load on the resting paw surface (Formozov, 1946). In winters of abundant snow and snow crusts tigers suffer from hunger and sometimes even die of starvation. In the southern Ussuri region on March 1 (14) at a time of crusted snow blood was noted on the tracks of a tiger. This animal was evidently very hungry ‘‘зшсе it stopped along the way to eat а half-pecked rotten wolf-fish washed ashore in the previous autumn.”’ Its feces “‘were small in quantity and contained . . . considerable undigested grass.’’ Some days later the carcass of the tiger was found in a nut pine Fig. 72. Nut pine forest at source of a stream—habitat of tiger. Sikhote-Alin preserve. Photograph by V.K. Abramov. 160 Fig. 73. Nut pine-spruce forest at the headwaters of the Ulakha— habitat of tiger. Here are also lynx, wild boar, wapiti and yellow-throated marten. Ussuri region. December, 1969. Photograph by A.G. Pankrat’ev. grove with no signs of wounds. The animal exhibited emaciation (Almazoy, 1890). In the Far East, after winters of abundant snow accompanied by high mortality among ungulates, the food situation sharply worsens for tiger. The winter of 1914/1915 in the Sikhote-Alin range was extremely snowy. Depth of snow cover reached 100 to 150 cm. In that winter wild boar mortality was very high and over the next three or four years very few were sighted; in addition, hunters killed most of the weakened wapiti. A sharp reduction in number of ungulates the following winter resulted in the starvation of tiger and lynx and a decline in their population (Kaplanov, 1948). Near Terneya in 1915 an adult male tiger killed was not only emaciated but its intestines empty. It weighed only 70 kg versus a normal weight of 200 kg (Formozov, 1946). This predator had inspected traps and devoured Siberian weasels caught in them, visited the camps of wintering hunters, and attempted to catch one of their dogs. In the snowy winter of 1935/1936, in the basin of the midcourse of the Iman, a young tiger weighing 110 kg, was found which had died of starvation. Its stomach was full of lichens (V. Abramov, 1962). 130 161 Fig. 74. Broad-leaved forests with nut pine and fir—habitat of tiger. Suputin preserve, southern Ussuri region. September, 1970. Photograph by V.G. Heptner. In the Sikhote-Alin tigers suffered from starvation in 1957 and 1958. In these mountains wild boar, the main prey of tigers, had almost disappeared. The snow cover had been very thick for two consecutive winters, 1.е., 1956/1957 and 1957/1958, depriving wild boar of their meager food due to crop failure of pine nuts and acorns. An epizootic plague further decimated the swine. At the same time, the wapiti population was reduced to about one-eighth its former level (K.G. Abramov). Starving predators began to wander extensively; attacks on sika confined in parks, and even on horses and dogs, were reported. Ten deer were killed in a short period in a deer farm at Gamov in the winter of 1959/1960. In the same winter at Kras- 130 Fig. 75. Coniferous forests along the middle Bikin — habitat of tiger. Ussuri region. Photograph by V.K. Abramov. norechka in the Chuguev region a female tiger broke into a courtyard and attacked a cow. In the Sikhote-Alin preserve tigers attacked horses on two occasions and several times stole dogs. Winter starvation has been a common phenomenon among tigers in Primor’e, since abundantly snowy winters are frequent there. Over the last 110 years severe winters have occurred in 1855-56, 1876-77, 1877-78, 1884-85, 1886-87, 1891-92, 1901-02, 1909-10, 1914-15, 1924-25, 1925-26, 1926-27, 1932-33, 1935-36, 1938-39, 1940-41, 1941-42, 1947-48, 1956-57, 1957-58, 1961-62, and 1964-65 (Nasimovich, 1950; Bromlei, 1963 and 1968). In the last 50 years, the winters of 1914-15, 1941-42, 1947-48, and 1957-58 were especially bad for ungulates and tigers. Such winters have also been recorded for the Pri-Amur (1887-88, 1941-42, and 1942—43).?8 In the Primor’e region winters with abundant snow occur т the winter of 1970-71 in the Sikhote-Alin preserve wild boar were very scarce, and `‘изег tracks were far more numerous than boar’’. Evidently the predator survived on Manchurian wapiti, which are more difficult for it to catch than wild boar. The tiger’s situation was eased by the fact that the winter was not snowy; they were not stressed (E.N. Matyushkin). In the southern Ussuri region conditions were such that tigers struggled to live and in that winter some attacked cattle and even broke into cattle sheds. (V.H.) 131 163 once in every two to nine years, and on the average once in every four years, sometimes consecutively for two to three years, and then tigers are especially hard hit. In the Primor’e exceptionally snowy winters recur once in six to fifteen years, on the average once in nine years. In such winters the snow cover reaches 70 cm in depth even by November instead of March, as is customary (Bromlei, 1963). Under unusual conditions in snowy winters, when an unbreakable ice crust is formed, hunting of ungulates by tigers is aided, since wild boar, wapiti, and roe deer slip while the predator moves well on such crusted ice. Tigers successfully preyed on ungulates in the Primor’e in the winter of 1877-78 because of ice crusts (Yankovskii, 1882). When moving through deep snow tigers quickly tire and often rest. Hence when the snow is deep or covered with a breakable crust, the predator sticks to rocky sites and sunny regions, and in moving from one area to another it often moves on the frozen bed of a river or stream, along wind-felled trees, pathways made by wapiti and other ungulates, or along man-made winter roads and ski tracks. If there is much snow on the ice, the animal tries to skirt it, choosing sections under coniferous trees where the snow is less. Fig. 76. Valley of Beloborodov spring with nut pine-broadleaved and nut pine forests along the slopes —site of tiger migrations along banks and valleys in the forest. Sikhote-Alin preserve. December, 1964. Photograph by E.N. Matyushkin. 164 The Amur tiger is quite well adapted to low temperatures; it has a luxuriant, dense pelage and fattens by autumn. Cubs have been recovered which had spent the entire night in the forest at temperature of —40°C, but without apparent ill effects. From time to time, however, animals have been found with both ears, and even the tail, frostbitten. Evidently the main factor limiting the spread of tigers northward is deep snow cover (over 30 cm) and, consequently, the absence of wild boar, wapiti, and roe deer. In Trans-Baikal and the middle Pri-Amur, tigers were distributed up to 52° М. lat., 1.е., farther north than at any other place in their range. The maximum depth of snow cover in this region does not exceed 20 cm and at some places is only 10 cm; its depth exceeds 30 cm only on the heights of the foothills (Rikhter, 1948). Food. In southeast Trans-Caucasus, the wild boar was the main prey of tiger. In the stomachs of tigers killed in the Lenkoran basin, only the remains of this ungulate were found (Satunin, 1914). Occasionally tigers preyed on roe deer, Caucasian red deer, and various domestic animals, including dogs; cattle were attacked only in winter (Dinnik, 1914; Vereshchagin, 1942). In Iran this predator preyed on the same species of animals, and also on goitered gazelle (Brandt, 1856). In Turkmenia, Uzbekistan, and Kazakhstan wild boar was likewise the primary food of this predator. Bukhara deer occupied second place in many regions of Middle Asia, and roe deer second place in Kazakhstan. In Tadzhikistan, Bukhara deer have held first place in the food of tiger in recent years, with wild boar and domesticated animals coming second. For example, in Tigrovaya Balka in the lower reaches of Vakhsh River, from 1946 through 1950 this predator attacked 19 deer, 17 cows, 4 horses, 5 asses, and 1 camel. In summer cows and horses straying from their herds or remaining alone at night became the prey of tigers in the tugais. The stomach of a tigress caught on June 26, 1950 in a tugai at Kyzylsu contained the meat and bones of cows (Chernyshev, 1958). In floodplain reeds tigers lie in wait for goitered gazelle which visit water holes; the remains of gazelles killed by tigers were once often encountered in the reeds (Flerov, 1935). In February, 1945 in Tigrovaya Balka the remains of goitered gazelle, wild boar, and deer were found in a den (Stroganov, 1959). In the lower Amu-Darya River tigers have sometimes killed and eaten jackals, chaus [jungle cats], and locusts (Pokrovskii, 1951). In Tadzhikistan, on the Vakhsh River, tiger feces often consisted of only locust remains, or an admixture of locusts and mole crickets, chitin of various beetles, hair of small mammals (mainly of complex-toothed rats [Nesokia indica]), and bird feathers (Flerov, 1935). In southern Tadzhikistan tiger feces sometimes contained the remains of mouse-like rodents, birds, and insects, and autumn and winter feces even contained kemels of oleaster (dzhida) and sea buckthorn fruits (Stroganov, 1961). 132 165 Fig. 77. Broad-leaved forest. Habitat of tiger. Southern Ussuri territory. Photograph by V.K. Abramov. On the Zhana-Darya and around the Aral Sea in Kazakhstan, in addition to wild boar, tiger also caught saiga, goitered gazelle, wild horses, kulan, and mountain sheep (Ovis ammon arcal) (Meyendorff, 1826; Eversmann, 1850). Goitered gazelles were preyed on during visits to water holes, and along the Ш (Alferaki, 1882). In reeds on the floodplains of the Syr-Darya, Chu, Ili, and other rivers, tigers evidently hunted saigas when they were numerous in these valleys during winter. In flooded areas and irrigation ditches tigers caught fish (sazan), which they eat with great enjoyment (Smimov, 1875). As in Tadzhikistan, in other regions of Middle Asia and Kazakhstan also, tigers attacked domestic animals, even camels, but most often dogs and horses. For example, in the winter of 1877 tigers killed all the dogs in a village beyond Chirchik at Кага Tyub (Smirnov, 1879). ‘‘At 132 166 places where cattle were abundant in Kirgizia tigers subsisted, especially in winter, almost exclusively on them’’ (Alferaki, 1882). In 1930 in southern Pri-Balkhash, in the region of Krasnyi town, tigers destroyed in a short period over 200 head of cattle, forcing the local populace in several places to migrate with their livestock to safer regions (N.Sh., 1930). Tigers formerly occupying Baikal and Trans-Baikal fed on wild boar, roe deer, wapiti, moose, and domestic animals, i.e., horses, cows, and reindeer. For example, the stomach of a tiger caught on November 22, 1953 near Gazinur River contained the meat and hair of a roe deer; another killed on December 24, 1953 in the Mogochinsk taiga contained the remains of a wild boar. ; In the Far East, in Sikhote-Alin and other regions, tigers live on wild boar, wapiti, moose, sika deer, roe deer, musk deer, Manchurian hare, bear (brown or white-breasted™), lynx, wolf, badger, hazel grouse, and even fish. Their primary food has always been wild boar in these areas. From the stomach contents of tigers and the remains of their prey, 11 animals have been identified over the years; of these, 3 were Manchurian wapiti, 4 wild boar, 1 moose, 1 musk deer, 1 bear, and 1 lynx (Kaplanov, 1948). Tiger food in these areas (expressed as a percentage of total stomachs and carcasses Fig. 78. Nut pine-spruce forest at headwaters of the Ulakha. Habitat of a female tiger with her litter. Wild boar, wapiti, lynx and yellow-throated marten also are found here. Sikhote-Alin. Deceinber, 1969. Photograph by А.С. Pankrat’ev. *Refers to Eurasian black bear (Ursus thibetanus), which possessed a large white patch on the throat and chest— Sci. Ed. [=] 167 135 examined) comprised: wild boar—40, wapiti—30, moose-—10, musk deer—10, bear—6, roe deer—3, and hazel grouse and other animals — 1 (G.F. Bromlei). In Sikhote-Alin preserve wild boar were found 21 times (35.7%), wapiti 13 (22.1%), moose 6 (10.1%), brown bear 5 (8.4%), musk deer 8 (13.6%), roe deer 2 (3.4%), lynx 1 (1.7%), and hazel grouse 3 (5.1%) (У. Abramov, 1962). In years when the population of wild boar was low and their physical condition poor due to compulsory feeding on horsetails and twigs, the role of different species of animals as tiger food changed. For example, from 1957 to 1959 the prey of tiger consisted of the following animals (based on number of carcasses found): wapiti—20 (50%), wild boar—12 (30%), brown bear—2 (5.0%), sika deer—2 (5.0%), and moose, roe deer, musk deer, and badger—1 each (2.5%). Evidently in years of poor yield of nut pine and oak, and winters of abundant snow, resulting in emaciation of wild boar, tigers attack them less often and hunt mostly for Manchurian wapiti and moose (K.G. Abramov; V. Abramov, 1962). From 1944 through 1950, 17 instances of tigers attacking bears (brown and white-breasted) were recorded. Old and sick tigers usually confine themselves to smaller prey: piglets, Fig. 79. Remains of bull wapiti devoured by a tiger. Beloborodov spring, Sikhote- Alin preserve. February, 1968. Photograph by E.N. Matyushkin. 137 168 young wapiti, roe deer, goral, and even hares and pheasants. For example, a lame tigress living in 1953 on the right bank of the Bir River, fed on hares which she caught on the island, and on wild boar piglets (Rodin, 1956). While prowling mountain streams and brooks, tigers skillfully catch fish by grabbing one in the water with a paw and throwing it on the bank. Turtles are scooped up when they surface for air. The infection of tigers by the helminth Paragonismus westermanii, the intermediate hosts of which are fresh-water crabs and crayfish, proves that tigers eat them also (V. Abramov, 1962). Tigers sometimes eat pine nuts, shell and all, wild berries, fruits, and summer grass. Carrion is resorted to only in time of starvation (Baikov, 1925) (Fig. 82).* In the last century, when tigers were numerous in the Far East, their attacks on domestic animals, especially dogs, were comparatively frequent. Such attacks increased toward the end of winter. In the 1950’s and 1960’s cattle killed by tigers became a rare occurrence there. From 1956 through 1959 in the Primor’e region, tigers attacked the following domestic animals: adult horses ten times, colts twice, cows ten times, and calves once. In the same period eight sika deer were killed in a deer farm (V.K. Abramov, 1962). Tiger attacks on domestic animals increase in years of a sharp reduction in population of wild ungulates, as occurred, for example, in 1957-1959. Apparently domestic animals near or in villages, or left to pasture without proper supervision, fall victim either to young predators around three-years-old or to old and sick animals which cannot hunt for their normal prey (К.В. Abramov). In our country tigers pose almost no threat to man; man-eaters, famous in India, have never been known here, although stray reports of attacks on man (see ‘‘Practical Significance’) have been noted. Man, however, has never been a “‘source of food’’ for Soviet tigers. In adjacent countries tigers feed mostly on large animals. In the northern part of northeastern China they hunt wild boar and wapiti, often attack dogs, and from time to time bear and small ungulates such as roe deer, goral, and catch hares and pheasants. The distribution of tigers in northeastern China (Manchuria) is generally associated with the availability of nut pines and wild boar: **Where there is nut pine, there is wild boar; where there is wild boar, tiger will also be found’’ (Baikov, 1925). In southern China wild boar, large deer, muntjak, domestic animals, and more rarely porcupines and pangolins serve as prey for tigers; tigers even eat frogs. In Yunnan in southwestern China their main prey are sambar deer, muntjak, and wild boar; on occasion they attack domestic livestock — pigs and buffalo calves (A.A. Sludskii). *Reference not clear; may mean Fig. 81—Sci. Ed. 136 169 Fig. 80. Young wapiti in velvet killed by a tiger. Ta-Chingou, Sudzukhin preserve, May, 1966. Photograph by У.Е. Prisyazhnyuk. In India tigers hunt for deer (sambar and axis), wild boar, buffaloes, nilgai, porcupines, peacocks, and domestic cattle; during floods they hunt for crocodiles, turtles, and fish. The stomach of some killed tigers were filled with locusts and mollusks. When starved, they catch frogs and mice (Jerdon, 1874*; Bikhner, 1905; Pocock, 1939; Tate, 1947; Berten, 1954*). Tigers rarely attack mature male boar. They avoid carrion but sometimes when food is scarce they will consume rotting carcasses, including those of other tigers. Sometimes tigers lick saline soil (Pocock, 1939). (Methods of stalking used by tigers are discussed under ‘‘Daily Activity and Behavior’’.) It has been stated that a hungry animal consumes some 32 to 48 kg of meat in one sitting (Baikov, 1925). This is hardly possible, although it has been established that a sambar may be consumed by a tiger in two days and a buffalo in three days, leaving parts of the prey unconsumed (Corbett, 1957). In one sitting a tiger reportedly consumed almost one-third (30 to 40 kg) of a killed wapiti. The meat of the croup was consumed entirely and the bones of the legs and vertebral column gnawed roughly up to the middle. *Not in Literature Cited— Sci. Ed. 170 The body of the марш was slit open across the chest cage. АП the organs above this line (lungs, heart, and liver) remained untouched while the intestines and part of the stomach were consumed (У.Е. Prisyaznnyuk). The normal daily ration of the tiger in zoological gardens is 10 to 12 kg meat (Obukhova and Shakhnazarov, 1949). In India, an adult tiger invariably kills no more than a single animal in one day. Young tigers, attacking a herd, may kill four or five cows in а single day (Jerdon, 1874*). In Sikhote-Alin an adult tiger kills and consumes per annum an average of 30 large animals of 100 kg (3,000 kg live weight). A family of tigers (mother and three cubs) consumed 280 kg meat in 20 days (wapiti 150 kg, wild boar 100 kg, piglets 20 kg, and musk deer 10 kg). In one year animals belonging to a single litter consumed about 5,040 kg meat (Kaplanov, 1948). In connection with the fact that tigers sometimes sustain prolonged hunger, in favorable seasons a large accumulation of fat reserves is found in the abdomen, groins, and body cavities in layers 4.0 to 5.0 cm thick. Fat layers are also found between the muscles. A young three-year-old male caught on February 16, 1940 in Sikhote-Alin yielded 30 kg fat (Kaplanov, 1948); a female caught on March 30, 1950 on the Pyandzha yielded 10.5 kg fat and the layer in her abdominal cavity was 40 to 50 mm thick (Chernyshev, 1958). Thanks to fat reserves constituting 20% of overall body weight, tigers which have eaten nothing for even five to ten days remain content. Evidently they can withstand even longer periods of starvation. Fig. 81. Brown bear killed and eaten by a tiger. Tatibe River, southern Ussun region. Autumn, 1961. Sketch by N.N. Kondakov from a photograph by V.E. Prisyazhnyuk. *Not in Literature Cited— Sci. Ed. 138 171 Tigers drink water often and in quantity. In Middle Asia, especially in winter, they mostly employ rivers as water sources, and not lakes. They drink from lakes mostly in spring and summer, when the brackish water is less saline than in winter. Home range. A diet of large animals compels tigers to prowl widely in search of prey and often follow ungulate from one pasture to another. Therefore, the dimensions of the hunting territory of an individual are very large. In the Ili delta, where wild boar and roe deer are numerous, one tiger covered a distance of roughly 50 cm x 20 cm along a stream and hunted periodically in different sections of this area. As soon as the number of pigs had been sharply depleted, the predator deserted the region and moved into another several hundred kilometers away from the former. In Tigrovaya Balka on the lower Vakhsh in Tadzhikisian, which abounds in unguiates, the territories of three tigers, though overlapping at places, covered an area roughly 6.0 to 7.0 km in diameter per animal (Flerov, 1935). In this same region, according to other data, the individual territory of a tiger usually does not exceed 30 to 40 km?.-However, tigers do not live for long at one place and in a few days or weeks move into an adjacent region or one farther away (Stroganov, 1961). In Sikhote-Alin preserve in the winter of 1939 to 1940 a single tigress periodically visited a whole system of rivers and ranges and was encountered in a territory 60 km x 70 km. An adult male, also in winter, was periodically sighted in a section 80 km x 40 km (Kaplanov, 1948). In this same preserve, in places rich in ungulates, tigers were sometimes confined to a territory around one or two springs covering an area of 400 to 500 km? (С.Е. Bromlei). In winters of heavy snow the territory regularly visited by tigers usually decreases, and the animal beats pathways along which it prowls. It may be assumed that tigers in summer spend a more settled way of life; yet even then they often move 10 to 30 km. A tiger prowls throughout its territory in a rather circular manner and hence is periodically sighted at the same place at intervals of a week or month, depending on the size of the animal’s domain. As in the case of other species of cats as well as bears, tigers demarcate their territory with characteristic markings; the animal rises on its hind legs and scratches the bark of trees with the claws of the forepaws (Baikov, 1915; Allen, 1938; Arsen’ev, 1950). The bark on a tree is sometimes stripped off to a height of 250 cm. From such markings the dimensions of the territory of a single animal can be judged. Special scent marks in the form of urine and excreta (tigers have preanal scent glands) are also deposited. For this purpose particular stones or trees are selected and periodically sprayed with urine. On finding a suitable site, the tiger sniffs it, then deposits its urine. Before undertaking a hunt a tiger will usually move along the top of 172 the mountain chain, from where it can inspect both slopes; often it rests on a high rocky outcrop overlooking a river valley and views the surround- ings. ‘‘Having killed and eaten its prey, either totally or partially, the animal roams its selected territory. At first it rolls in the snow every 100 to 200 m, leaving spots of blood in the print of its body, which reveal its departure from a recent kill. During the day the animal often rests on its abdomen, urinates, defecates, and paws the snow and ground quite often, moving tens or even more kilometers before lying down in a clear sunny section devoid of trees. At night, on the contrary, it selects for rest a dense spruce thicket which ensures a few degrees more warmth than a sparse growth’’ (Kaplanov, 1948). After resting, which may continue for 12 or more hours, the animal resumes its journey, only rarely stopping to roll in the snow. During long- distance forays, periodically stopping and resting, sometimes however without stopping, a tiger may travel for several days and cover some tens, even hundreds of kilometers. In a single day the animal may traverse 20-50 (Kaplanov, 1948) to 100 km (Baikov, 1925). Only during estrus do the female and male stay together in a comparatively small area. Tigers prefer an area rich in wild boar or Manchurian wapiti. Unlike single animals, a tigress with cubs less than two years old will occupy a very small territory. For example, in Sikhote-Alin, in a section rich in ungulates, a tigress with cubs lived in an area 5.0 km x 3.0 km from December 24, 1940 through January 15, 1941 (Kaplanov, 1948). A tigress with cubs aged 2.0 to 3.0 years will extend her territory much beyond that. In 1957 in the upper reaches of the Maikhe River a family consisting of a tigress and two cubs in their second year was discovered. Until the spring of 1958 this family lived in a fairly limited territory. Later they enlarged the zone of their activity and by autumn of 1959 occupied an area of 50 km x 60 km (V. Abramov, 1962). Burrows and shelters. Solitary tigers usually do not establish a perma- nent den but Пе down to rest close to their kill. In the river vaileys of Middle Asia and Kazakhstan favored sites for rest during the day are the edges of extensive reed beds adjoining a tugai or open forest glade. The den is usually situated in the shade of a single oleaster (dzhida) or turanga tree [Euphrates poplar] occurring among reeds or reed grass [veinika], which becomes strongly trampled down. Dry grass serves as a bed. At times a very distinct path to the den is apparent in trampled grass, adjacent to an area 30 m x 40 m? of trampled or beaten grass scattered with bones, bits of skins, and wool discarded by the animals. Obviously the stench from such areas is hardly pleasant. Judging from trampled grass the animals undoubtedly rest in several places, moving with the shade (Stroganov, 1961). In the Far East tigers select for rest a fairly concealed place somewhere under a nut pine, sometimes even on sunny sands or rock. They return fairly ВУ» regularly to favored sites and make a den. In winter they rest directly on the snow. A tigress hides her cubs in rocky areas of the mountains or in dense vegetation, selecting a niche in the rocks, cracks in a precipice, or a cave for a den. ) In Middle Asia and Kazakhstan dens for cubs are located in dense galleries of reed or in tugai, on an islet unaffected by water. The den is established in a natural soil depression and sometimes covered with dry leaves and grass, but usually lacks bedding. Often the den is well concealed by dense reeds, wild vines, lianas, and other plants. A tigress with very young cubs will approach her den cautiously and attempt to leave behind as few traces as possible. Dens are established in areas of abundant wild boar and other ungulates. A tigress uses the same den for several consecutive years. The den of a dead tigress is often taken over by another female. When the cubs grow up and begin to follow their mother, she leaves them behind in a temporary den nearby, within 500 m, when going out to hunt. Hunting is done stealthily so as not to frighten ungulates wandering close to the den (Kaplanov, 1948). In India tigers set up dens under rocks or in caves. On one occasion, however, a pair of cubs was found beneath a prickly bush (Jerdon, 1874; Pocock, 1939). Around the den scattered bones and excrements often exude a characteristic strong odor. However, within and around the den per se, as long as her cubs are small, the mother leaves neither bones nor other remains of prey (Anderson, 1964).* Daily activity and behavior. Within the Soviet Union tigers become active around twilight; they may hunt at any time of the day but most often after sunset, in the first half of the night, or later at dawn. Tigers cannot withstand heat well. In India they usually set out to hunt at sunset and hunt throughout the night, slowly covering their territory along trails (Pocock, 1939). Tigers employ several methods for hunting wild boar, wapiti, moose, and other animals, but mainly resort to ambush. On finding the track of a wild boar or wapiti, the predator follows the spoor until it comes fairly close to its prey. Here it turns to the leeward side and concealing itself on the path awaits the approach of its intended victim. The olfactory sense is rather poor in tiger but nevertheless the animal is capable of tracking a quarry even when the spoor is an hour old. The predator locates its quarry mainly through the senses of audition and vision. Hunting tigers are assisted by the bright and variegated color of their coat, which blends with the surroundings. When a tiger runs through the forest among shrubs the black, yellow, and white colors of its coat merge and the animal appears a monochromatic brownish- gray (Arsen’ev, 1949). When hunting in tall reed thickets tigers sometimes *Not in Literature Cited — Sci. Ed. 140 174 rear up on their hind legs or leap upward in order to inspect the surroundings (Khakhlov, 1928). In summer tigers lie in wait for wapiti, moose, and roe deer at back- waters, where these ungulates go for food, and at ponds, and also at salt licks; in winters of heavy snow tigers maintain a vigil along their trails. The predator approaches the quarry closely in one or two bounds and throws it to the ground by striking it with its forelegs. The toppled prey dies almost immediately due to cervical vertebrae broken at the occiput or a broken occiput per se. A wapiti killed in Sidzukhin preserve had a broken neck; on the right shoulder, struck by the tiger’s forepaws, the skin was torn and showed the imprint of all five paw cushions, the same as a pug mark in mud (V.E. Prisyazhnyuk). Huge canines help the tiger kill an animal quickly. It requires more time only for large wild boars, bears, and bulls. When dealing with big game tigers attack from the sides and rear. Having felled its prey the predator does not catch hold of it immediately but makes a few more strikes or pounces. A tiger will give up the chase after pursuing a potential quarry for about 100 to 200 m. Tigers have adopted a unique method of hunting in the Ta-Chingou area in the Sudzukhin preserve (Figs. 71, 80 and 84). There wapiti and sika deer regularly wander onto the flat sandy shores of the bay at night and sometimes even in the day. On the night of Мау 23* the fog was heavy. Taking advantage of this fact a tiger approached within 10 to 12 m of a herd of seven wapitis standing right at the edge of the water. Launching its attack from behind a small undulation, the predator overtook one of the velvet antlered wapitis [Fig. 80] in three short bounds. Later, it dragged the carcass half-a-kilometer into a grove of hazel and young oak trees. Hunting by the sea in fog is the usual method employed by tigers in this region. The fog is so thick that a human is not visible until approached to 20 to 30 m. Tiger tracks in such mists on coastal sands right at the water’s edge sometimes cover a kilometer. Spoor are visible there both in summer and winter (V.E. Prisyazhnyuk). Having tracked an animal, a tiger will not run long after its quarry. The only exception is the wild boar, which a tiger chases with tenacity; the predator will systematically kill one after another until the entire herd is destroyed. Subsequently, it will set off to find a new herd. Hunters refer to this method of tiger hunting as ‘‘grazing’’ the pigs. In attacking animals a tiger makes very little noise and hence does not scatter a herd. In Sikhote-Alin preserve a single tigress stole one animal after another from a herd of wild boar, destroying the lot in just a few days. In spite of repeated attacks boars still remained in the same section of the preserve. Shortly after the tigress, a male sneaked up on another family of *Year omitted in Russian original —General Editor. 141 175 wild boars, made a couple of pounces, and catching one of the pigs carried it in bounds 200 m away into a bush. Having consumed the catch, it repeated this tactic a couple of days later. The pigs in the herd, though uneasy at night, appeared calm and grazed quietly during the day on the sunny slopes of the river. On another occasion an attack by a tigress on a resting female wapiti was observed. The attack was so swift that the deer had no time to even stir from its bed. In December, 1941—January, 1942, working at the site of permanent tiger occupation in the basin of,the Fata and Sitsa rivers (Terneya region), G.F. Bromlei (1964) did not sight even once the exodus of wild boar due to their decimation by this predator. Yet he found the remains of slaughtered pigs in several places. From the traces it was established that the tiger hunted at night. Sneaking to within 10 to 12 m of a herd of pigs, it executed two or three bounds, caught hold of the victim, and carried it away without shedding so much as one drop of blood. Only from a narrow trail on the snow, caused by the dragging legs of the prey, was Bromlei able to establish that the predator had succeeded in its hunt. The remaining boars, frightened by the attack, scattered from 200 to 300 m and then settled into a fresh shelter for the night. In the Soviet Union this predator is relatively silent during a hunt. Finding himself during the course of one month in a region where tigers jive, G.F. Bromlei never heard a tiger voice even once. In India, contrarily, during a hunt and after it, a tiger roars frequently (Corbett, 1957). A tiger will even tackle a bear, sometimes one much larger than itself. Having tracked its victim, the predator selects a rock or windfelled tree from which it can ambush the quarry from the leeward side. It springs upon the approaching bear from an overhead position, catches it by the chin with the claws of one forepaw and the throat with the claws of the other, and clamps its jaws on the spine. Bears are generally afraid of tigers and, coming across their tracks, run away perpendicular to the trail. Confronted with a tiger, a bear will try to escape attack by climbing a tree (Baikov, 1925). On February 6 in Sikhote-Alin a tigress discovered the den of a female bear with yearlings. Digging up the den from the side opposite to the entrance, she proceeded to harass the mother from both sides alternately. Eventually the tigress seized the bear from the rear with her claws, behind the legs, dragged it out of the den, and quickly killed her by clamping its jaws around the cervical vertebrae at the occiput. The cubs, weighing 30 kg, each, she killed within the den (Kaplanov, 1948). In the Ussuri territory the hair of bears has been found repeatedly in the spring and autumn feces of tigers (April 24, May 7, and November 10, 15, and 27). Early in May, 1951 on the bank of Tatibe River (Iman tributary) a bear was found (body length 158 cm, weight 170 kg), which had obviously been mauled by a tigress 148 176 (Fig. 81). АП the fatty parts of the body had been devoured—the back, hams, and fatty layers in the groin region. Fecal deposits, urine trails, and three separate lairs within 10 m of the carcass of the bear in which the tigress had laid up for three or four days were found. In the upper reaches of Pfusung River (west of Ol’ga Bay, Primor’e region) in September, a small white-breasted [Eurasian black] bear climbed up and down a tree several times trying to save itself from a tiger. Evidently, the tiger harassed the bear a long time since the bark of the tree was totally peeled off at places. In 1949 at the end of December, in the basin of the lower Sinanche (Iman tributary), hunters chasing a family of tigers found a white-breasted bear hiding from one in a tree. Judging from its badly Fig. 82. Track of a tiger (male) traversing deep snow with impression of its tail also visible. Beloborodov spring in Sikhote-Alin preserve (Sitsa basin). February, 1968. Photograph by D.N. Matyushkin. 142 ТАТУ mauled skin and tufts of detached fur and other traces оп the snow, the bear had been wounded by the tiger’s claws and remained in the tree for over a day. Brown bears fall victim to tigers more often, since white-breasted bears usually live in hollows or among close-set rocks and are thus less accessible. In the middle Sikhote-Alin only three instances of killing of white-breasted bear by tigers have been recorded over several years of observation. Over 15 instances of tiger attacks on brown bears resting in dens located on the slopes of the middle Sikhote-Alin (Sikhote-Alin preserve) were recorded from 1952 to 1959, mainly in late autumn and early spring. Tigers did not always succeed in killing their quarry. The largest of the bears escaped from the tiger’s claws, and having been chased from their dens, began to wander in the taiga, becoming ‘‘тоуегз””. From 1940 the number of tiger attacks on bears has risen steadily, due, no doubt, to a general reduction in ungulate population. Evidently in the absence of ungulates tigers resort to feeding on bears (Bromlei, 1965). As mentioned above, a tiger is not always the victor in an encounter with a bear. In the winter of 1959/1960 a tigress lived with her two-year- old cubs in a ravine in the Sikhote-Alin. Once, having killed a wild boar, she left it with her cubs and went away. A large bear wandered into the area and on her return to the den the tigress attacked it. During the ensuing contest the bear killed the tigress. In spite of serious wounds, the bear proceeded to eat the wild boar and then the tigress (Sysoev, 1960). In 1913 on the Bol’shoi Sinanche River a large brown bear knocked down a tiger. In 1960 in Sikhote-Alin preserve a young tiger became the prey of a bear (V. Abramov, 1962). A similar incident has been reported in the Pri-Amur (Krivopusk, 1957; Fig. 88). A hungry male or a lone tigress, after killing a large animal weighing 100 to 150 kg, eats heavily and rests in the vicinity for five to ten days. Usually a tiger remains for only three or four days by its prey, until the meat begins to freeze solid or to decompose. Sometimes, having killed an animal, the tiger leaves it untouched and returns to it later. In such an event it sometimes covers the kill with brush, snow, and grass which it uproots with its teeth. It consumes large prey skin and all, leaving only the feet, hooves, and head. It even cracks the long bones of moose. Having killed an animal the tiger usually drags or carries it into a bush close to water. After feeding it rests for much of the day, lying on its side with the legs outstretched, getting up occasionally to drink from the water hole. If hunting has been unsuccessful, it may continue to hunt even during the day. Like other species of cats, tigers are very clean animals. They often lick their coat and paws. After a hearty meal they love to roll on the snow 178 and in summer bathe in a river, submerging the muzzle in water and ‘‘gargling’’ to rinse the mouth free of remnants of meat and blood. Claws are cleaned by stripping soft bark. On hot days, as a defense against blood- sucking flies, they lie in water with only their head protruding, or stretch out in the spray of a waterfall. Tiger movements are graceful, quick, and nimble. They can creep through grass only 60—70 cm high and remain completely undetected. The animal’s pace is usually 50-80, and up to 100 cm long; it begins to sprint only when chasing its prey and may then leap as far as 4.0 m. The maximum length of a leap is 6.0 to 7.0 т, and a height of 2.0 to 3.0 т. Tigers negotiate precipices and rocks very well but generally cannot climb a vertical tree unless it has close-set branches. Exceptions are known, however. A tigress pursued by hunters clawed up a tree devoid of branches to a height of 7.5 m (Pocock, 1939). Tigers are excellent swimmers, and large rivers (Amu- Darya and Amur) or sea channels up to 5.0 miles wide pose no problem. Adult males and single females not in estrus live alone. A tigress with growing cubs aged 1.0 to 3.0 years moves with her offspring; the latter sometimes comprise cubs from two litters. This explains why tiger families of four or five animals are sometimes seen. When tigers were abundant in the Far East, groups of even 7 to 13 animals were recorded (Kaplanov, 1948). If such groups actually existed at all they probably represented two families which for some reason had joined together. The tiger possesses tremendous strength and is capable of carrying off animals far larger and heavier than itself such as horses, cows, and buffaloes up to three years of age (Brandt, 1856). A tiger can transport a horse or cow tens and even hundreds of meters from the place where it killed, it being carried, not dragged. Corbett (1957) tracked a tiger which carried a fully grown cow 4.0 miles (7.5 km) away from the place of its kill. Another tiger dragged a killed bull 13.5 meters. The bull could not be moved by 13 men. A tigress was seen swimming across a river holding a cow in her mouth (Pocock, 1939). Once a tiger jumped into a pen, killed a calf, and catching it in its teeth leaped back to a rock situated 3.6 m above the pen’s level (Allen, 1938). If the prey is so heavy that the tiger cannot carry it, it is often abandoned. When the predator carries its prey by the neck, the rear of the body drags, leaving behind a distinct trail. If, however (as for example, a deer), the tiger holds the kill by the middle of the back, the portage trail may be faint or nonexistent. A tiger can carry a large wild boar in its teeth through dense rushes, jump onto rocks, and even climb a steep slope. This predator is less afraid of humans than other large animals, but nevertheless usually moves away from them. Tigers usually initiate an assault only when injured or chased, confronted by dogs, being approached at a den with cubs, or in an unexpected or chance encounter at night. Tigers are 143 NAS) generally more aggressive at night and approach man more boldly (Kaplanov, 1948). A tigress will sometimes attack to protect her cubs. Ussurian hunters who had captured several dozen cubs reported only one instance. when a tigress, frightened off by shots fired in the air, returned to her litter and had to be killed. A tiger does not attack when men approach its kill (also see section ““Territory’’ for further information on the behavior of tigers). Tigers caught young respond well to training. Even adult animals can be trained. Seasonal migrations and transgressions. It was noted above that single tigers wander much of the time in search of prey in a definite and fairly large region. Sometimes these wanderings are regular. For example, in Kazakhstan and the Far East tigers in spring climb high into the mountains, following grazing ungulates, and descend to lower altitudes in autumn. Periodic emigrations of tigers are definitely associated with a reduction in population of wild boar, or their migrations, or fires in the taiga, and in large sections of floodplains, and tugais. In Tadzhikistan it has been noted that an increase in the tiger population is directly related to massive fires in the tugais on the Afghan banks of the Pyandzh. In the lower reaches of the Syr-Darya tigers have been sighted only twice in the last 30 years, once in 1926 and again in 1945, and on both occasions were trailing herds of wild boar moving along the coast of the Aral Sea from the delta of the Amu-Darya. On the latter occasion they covered over 1,000 km calculated in a straight line. Reportedly, tigers following wandering wild boar in winter passed through the Kyzylkum. In 1930 in the Ш delta, consequent to large-scale slaughter of wild pig for meat, the tiger population dropped sharply. As a result, tigers emigrated from this area that year. Two moved upstream along the river: one was killed 500 km from the delta and the other entered China along the valley. Tigers caught in the last century in central Kazakhstan and also at Biisk and Barnaul were intruders who had left their permanent dwelling areas (Balkhash and Zaisan) located at a distance of 400 to 600 km in a straight Ппе.” In southeastern Trans-Baikal in the last century tigers were sighted only periodically, once every five to eight years. At that time they were killed near the Argun, Gazimur, and Shilka rivers. Following reindeer, the predators intruded into Stanovoi range from the upper reaches of Amur and Shilka (Brandt, 1856; Sel’skii, 1856). In southeastern Trans-Baikal they were seen from time to time early in the present century as well as in the 1950’s and 1960's. In 1953 about eight tigers were reportedly seen there (А.М. Leont’ev). One was killed on November 22, 1953 at Gazimur River in the Ust’-Kara Recognizing this habit of tigers—to wander or undertake extensive migrations, Kazakhs popularly call them ‘‘road’’ or *‘traveling’’ leopards. 144 180 region of Chita district, and another in the same year on December 24 оп. the bank of Shilka in the Mogochinsk region (Sludskii, 1966). From 1963—1966 tiger tracks were detected repeatedly along the Uryumkansk River in the Plyusnino region and along the Gazimur River near Kungir, Kurumdyukon, Kaktolg, and Kachugai stations (Sinitsyn, 1966). Evidently they entered Trans-Baikal from the Great Khingan mountain range in China or from the Amur region. A tiger caught in 1905 close to Ust’-Maya (Aldan; Ognev, 1935; Galchenko, 1959) had traveled 1,000 km in a straight line from its region of permanent residence. Animals seen from time to time in the Verkhoyansk range (Silant’ev, 1919) and crossing the Stanovoi range up to 56° М. lat. had traveled no less distance. As mentioned above, they used to move that far north in pursuit of migrating wild reindeer. In the Far East, around 1886, tiger migrations were recorded for two consecutive winters moving west to east, from the Sungari in China to Shikote-Alin. All of the tiger tracks ran eastward according to Arsen’ev (1949). A similar migration was observed in 1913 (Kaplanov, 1948). There are records of annual migrations of tigers from northeastern China (Manchuria) into the Soviet Union, the region of Bikin River and other places. In 1924 hunters came across the track of a large tiger and tracked it. The chase continued for 22 days and in that period the tiger had moved from the Tetyukha to Taukha rivers, covering a distance of 1,000 km. Later the tiger crossed into the Suchan valley and from there to the Daubikhe River, where its trail was again picked up by the hunters, who followed it for another 15 days up to the Iman (V. Abramov, 1962). In earlier years tigers intruded almost every autumn for a few days into the Suputin preserve on the Suputin River (southern Ussuri region), descending from the Dodyan’-Shan mountain to the sources of Kamenka and Anikinsk springs. At other times of the year their tracks were never seen in the Suputin preserve (Bromlei and Gutinkova, 1955). In the last century tigers were reportedly seen on Sakhalin [Island] in the Tym’ -Poronai valley in the upper reaches of the Tym’ and in the mountains near the Poronai River (Schrenk, 1858; Shmidt, 1868; Przheval’skii, 1870; Chekhov, 1956). Information is available, which requires confirmation, that tracks of a tiger were encountered on that island in 1950 and 1965. Tigers could have reached Sakhalin by swimming the 7.0 km width of Nevel’sk Strait. Because of the very deep (3.0-4.0 m) and porous snow cover, a permanent residence for tigers on Sakhalin Island is impossible (for other tiger transgressions see ‘“‘Geographic Distribution’’). Tiger migrations into the Soviet Union from the Korean Peninsula have also been observed. At the end of the 1890’s tigers crossed on ice over the 181 Amlokan’ River into the Korean Peninsula from northeastern China (Manchuria) (Von Hon Gu). Reproduction. Estrus, gestation, and parturition among tigers of northern subspecies, according to the latest observations in the Far East and in zoolo- gical gardens, are not restricted to any particular season. Newborn cubs may be found at any time of year. For example, in December, 1932, a month-old cub weighing about 2.5 kg was caught in the Ussuri region on the Baitsuka River, the right tributary of the Iman. In January, 1933, four dead cubs weighing 4.0 to 6.5 kg were found on the Iman River. In May, 1933 a female tiger was caught near the Sanchekheza River with five fully developed fetuses in her womb. In the winter of 1937/1938 three cubs aged three or four months were found on the upper reaches of the Iman (Salmin, 1940). Small cubs were found in this same region on January 9, April 13, July 22, October 19, and December 7 and 13 (С.Е. Bromlei). In March, 1957, a litter of four cubs aged 1.0 to 1.5 months was found (K.G. Abramov). A female caught on March 30, 1950, in southwestern Tadzhikistan at Pyandzh was in heat and had recently mated with a male as detected from spermatozoa present in the swab of the uterine cervix. A second female, caught along the Kyzylsu on June 26, 1950, was lactating (Chernyshev, 1958). Cubs have been sighted in Kazakhstan from June to August (N.A. Severtsov, 1861). Early in February, 1909 feces of four cubs roughly one- month old were found on the lower reaches of the Ili. In early April, 1908, on the shore of [Lake] Balkhash in the region of the lower Ayaguz three small cubs were caught (Shnitnikov, 1936). In Soviet zoological gardens tigers in heat can be seen at any time of year, once every two or three months, but sometimes every month, or only twice a year. Three Ussuri tigresses in the Moscow Zoological Garden gave birth as follows: one (“‘Chizhik’’) twice in March, once in April, twice in May, and once each in October and November; another (‘‘Naya’’) once each in May and June; and the third (‘‘Lyuba’’) twice in May (Afonskaya and Krumina, 1956). Births in zoological gardens occur most often in two seasons—spring and late autumn; however cubs usually appear in May (46% of all newborns). Thus the periods of heat formerly reported for different regions of the USSR (January and February in Trans-Baikal, December and January in Kazakhstan, and end of December and January in the Far East) are only partly correct. In the northeastern part of China newborn cubs are usually seen in April to June but most often in May. In southern China small cubs are seen not only in spring and summer but also in winter. In southwestern China (Symao, Yunnan) cubs are more often seen in April to June and September to October. That Indian tigers have no definite breeding season has long been known (Jerdon, 1874). The absence of strict seasonal periodicity in 145 182 the appearance of litters among tigers points to their southern origin. Nonseasonal reproduction is characteristic of many tropical animals. Estrus among tigresses continues for 12 to 18 or, rarely, 25 days. Among Ussuri tigresses in the Moscow Zoological Garden, it extends for 3 to 13 days (Afonskaya and Krumina, 1956). A female in heat makes a charac- teristic rumble, sniffs frequently, micturates often, rolls on her back, splays her legs, and plays with the male. She eats little or no food. During this period she usually moves with a single male, but in areas where tigers are numerous, two to six males follow a single female. During heat tigers roar more often than at any other time. The female attracts males by her roar. Serious fights occur among males. They also become less wary during rut. The mating period of tigers extends for 5 to 8 days, and sometimes up to 18. According to observations made in the Moscow Zoological Garden, up to 20 matings occur in a day, with an average of 11, during the mating season (at night the animals separate). The interval between successive matings may vary from three to four minutes to several hours. In all, 43 to 123 matings have occurred in this period. While mating, the tigress lies on her chest and elevates her rear; the male grips the nape of her neck in his teeth (Afonskaya and Krumina, 1956). According to observations in zoological gardens, the duration of gestation is 98 to 112 days (Satunin, 1915; Baikov, 1925; Pocock, 1939; Shereshevskii, 1940). Commonly, gestation requires 105 days. Among Ussurian tigers gestation extends for 95 to 107 days, with an average of 103 (calculated from the first to the last mating). At the end of estrus the female isolates herself from the male. In zoological gardens after the mating season males and females have to be separated immediately since fights ensue between them. The male does not take part in raising the young. Tiger litters throughout the USSR consist of two to four cubs, rarely one, and even more rarely five or six. Cubs die quite often in the first few months due to various causes and hence two or three are usually seen with the mother and more rarely one or four. In different parts of the range in the Soviet Union the following number of cubs have been recorded in a litter; Trans- Caucasus — мо litters with two cubs each; Kazakhstan—three with two each, one with three, and one with four; Tadzhikistan—two with one each; and in the Far East—seven with one each, seven with two each; eight with three each, and two with four each. Additionally, a female with five fetuses was caught in the Far East. I.T. Trofimova, the tiger trapper, came across a female with four two-year-old cubs in 1953 in Sikhote-Alin near the Tudo-Vak River. From 1936 through 1957 hunters found five litters with two cubs each, ten with three each, and two with four each. From 1957 to 1959, four litters were registered with one cub each, eight with two each, and three with three each (V.K. Abramov, 1962). Among Ussuri tigers in 146 183 the Moscow Zoological Garden the number of cubs in a litter is the same as in the wild. Most often two cubs occur in each litter (50% of all cases) but the fertility of various females is not identical. The tigress “‘Chizhik’’ usually gave birth to three cubs (Afonskaya and Krumina, 1956). In the Peking Zoological Garden in 1958 there were either two or three cubs in the litters of two Ussurian tigresses. In the.southern part of the range, for example, in the South China tiger in Yunnan province usually one or two cubs occur, rarely three or more, although a case is known here of a female captured with six embryos (A.A. Sludskii). Other southern China tigers produced two cubs (twice) and four cubs (twice) (Allen, 1938). In India two cubs are common, more rarely three or four, and very rarely five or six (Jerdon, 1874; Pocock, 1939). Tigresses with seven embryos have also been reported (Beriff, 1932).* Growth, development, and molt. Cubs are usually born blind and helpless but sometimes with their eyes open (evidently when gestation is greatly prolonged). The weight of newborn Ussuri cubs varies from 785 to 1,043 g, body length 31.5 to 40 cm, tail length 13 to 16 cm, and length of ears 1.5 to 2.5 cm. The claws are not pigmented. Cubs born blind gain sight on the sixth to the eighth day (according to other data on the fifth to the tenth day). The ears open on the fourth to fifth day and the young begin to respond to sounds at some distance by the twelfth to the fifteenth day. There are no teeth at birth but the places where they will appear are distinct on the gums. Cubs cut the two medial incisors on the 13th day. Two incisors are seen on both the upper and lower jaws on the 23rd day, six incisors on both the upper and lower jaws on the 42nd day, and upper and lower canines on the 53rd day. There are two cheek teeth, two canines, and six incisors on the upper jaw on the 63rd day. At the age of 8.5 months the cub sheds the middle pair of incisors and the other incisors at the age of 9.0 months. Molt of the juvenile fur coat occurs at the age of 3.5 to 5.5 months (Afonskaya and Krumina, 1956). The color of the juvenile coat is the same as that of adults but the background much lighter and the stripes light brown. The ears of small cubs are relatively large. As in adults, a bright black color with a white spot in the middle occurs on the back of each ear. Therefore, this signal color is fully developed at an early age. Cubs grow and develop rapidly. On the 12th to the 15th day they begin to crawl around the den and by the 20th to the 30th day emerge, walk on the snow, and climb trees. Even two-year-old cubs weighing up to 60 kg may climb trees, but most lose this ability early. The growth rate of young tigers can be judged from the weight chart of ‘‘Sirotka’’, a cub fed artificially in the Moscow Zoological Garden: *Not in Literature Cited — Sci. Ed. 184 Age Weight Age Weight (days) (kg) (days) (kg) oh 0.8 133 18.7 31 2.0 180 21.8 63 4.3 210 26.5 93 10.0 284 38.5 When 35 to 36 days old cubs begin to lick meat and in 43 days attempt to eat it but receive no encouragement from their mother. After 48 days cubs tear off bits of meat. At the age of two months they begin to eat meat regularly but the mother continues to suckle them to the age of five or six months. The mother transports two-month-old cubs from one den to another. Leaving them there she sets off to hunt, and having caught a prey, takes her offspring to it. The litter remains near the kill while she again sets out to hunt. As long as the cubs are small the mother leads them to each kill. She does not leave her offspring of less than six months of age alone for long; those older than that she leaves alone for five or six days and two- year-olds for up to fourteen. Very young cubs left alone in winter sometimes freeze to death (Salmin, 1940; G.F. Bromlei). When prey is plentiful the litter is not moved far and remains in a relatively small area. Cubs are extremely agile and playful. When their hunger has been appeased they romp continuously and play with their mother until they feel sleepy. In play one cub will hobble side-ways, hide pretentiously, and wait for another cub to discover it; it then jumps out and tries to pin the other down by catching hold of its hind legs or tail. Cubs play with the mother in the same way. Sometimes a cub having caught another by its tail with its teeth follows behind the captive for quite some time, refusing to relinquish its prize. Cubs seldom fight or quarrel. The young scampering around a prey trample large areas and bite off branches from tree and shrubs nearby. If prey is scarce in the locality of the young and their mother absent for a long time, hungry cubs will gnaw at rotten wood. Cubs remain with their mother for two, sometimes even three years, and begin to live independently only in their fourth year. This biological feature of tigers has been remarked upon before. ‘‘It has been said that old babry feed their cubs for three years, during which period the latter are so quiescent that they can be caught without fear of attack’’ (Rychkov, 1762). In 1953 on Sikhote-Alin, I.T. Trofimova, the tiger catcher, came across a family of three three-year-olds and caught them all without much difficulty. By two years of age young tigers weigh over 100 kg. In the winter of 1950/1951 in the Ussuri region ‘‘lonchaki’’ (in their second year) [1.е., yearlings] weighing 113 kg each were caught (K.G. Abramov). A three- year-old male sometimes weighs up to 150 kg. Young tiger of this size 147 185 generally hunt with their mother. From time to time cubs from two different litters have been sighted with a tigress, one or two being cubs from an earlier litter. Males and females usually attain sexual maturity by their fourth year. In the Moscow Zoological Garden three female Ussurian tigers attained sexual maturity between the ages of three years four months and three years eight months. One male was sexually mature at the age of three years eight months; he attempted mating, however, at the age of two years seven months (Afonskaya and Krumina, 1956). Since cubs remain with their mother for a long time, she only litters once in two to four years. It is known that ““babry ... Whelp once in three years’’ (Rychkov, 1762). In the Moscow Zoological Garden, however, one female whelped twice in the same year and became pregnant a third time (Salmin, 1940). This female did not suckle her cubs. During the course of her lifetime a tigress generally produces 20 to 30 cubs of which usually one-half perish within six months (Berten, 1954*). In the Soviet Union a tigress produces only 10 to 15 cubs (A.A. Sludskii). According to observations made in captivity a tigress retains her ability to reproduce up to 20 years of age and her longevity is 40 to 50 years (Baikov, 1925). The Ussurian tigress Sirotka born in the Moscow Zoological Garden lived for only 18 years. In India one tiger survived in a relatively small region for 15 years but was killed in its prime. Another predator attacked cattle of a particular region for 20 years and, when killed, was not at all senile (Pocock, 1939). Tigers molt twice a year; in the Ussuri territory molt takes place in September and October and again in March and April. The long and sharp claws are shed annually in late autumn. At the time of shedding their claws, tigers often scratch the soft bark of cedar, Amur cork, and fir. Enemies, diseases, parasites, mortality, and competitors. The adult tiger has no enemy in the animal world. Very rare indeed are instances of its death due to injuries inflicted by a male boar, buffalo, or bear. Tiger mortalities due to wild boar have been recorded in the Trans-Caucasus and Iran (Brandt, 1856), Kazakhstan, Middle Asia, the Far East and India. In the upper reaches of the Iman in the Ussuri region a wild boar gored a tigress to death, and her two surviving cubs died of starvation. In 1933 in the Baikin region a tiger weighing 120 kg was killed by a boar. Near the Bol’shoi Sinanche River in 1913 a large brown bear crushed a tiger (V. Abramov, 1962). Cubs are killed more often than adult tigers. They are killed by male tigers, brown bears, and other predators. This was especially apparent in 1956 when the yield of pine nuts and acorns in Birobidzhan was inadequate *Not in Literature Cited — Sci. Ed. 148 186 and wild boar could hardly find food. Many brown bears wandered late in autumn because of the scarcity of food, instead of remaining in their lairs. One such bear on the Shukhi-Pokto range in the Khingan taiga attacked a three-year-old tiger and tore it to bits. Judging from the marks on the snow, the sequence of events may have been as follows. A tigress moving with a large cub attacked a herd of wild boar and wounded a pig. Leaving her cub with the quarry, the mother went away. A roaming bear came across the young tiger and the prey and a fight ensued between the predators. Tufts of fur from both the animals and numerous blood splotches were found scattered on the snow. The bear won. Having carried the pig to a wind- felled tree, the victor ate from it several times. After the boar was consumed, the bear returned to the tiger but was frightened off it by people (Krivopusk, 1957). Another instance of a wandering bear killing an adult tigress in Sikhote-Alin has been described above. In 1960 a young tiger was killed in combat with a bear in the Sikhote-Alin preserve (V. Abramov, 1962). In India tigers suffer greatly and sometimes even die from porcupine quills, which stick in their body when the rodent is attacked. A man-eater that was shot had 102 porcupine quills extracted from it. Some of these quills were 20 cm long and as thick as a pencil. Most were embedded in the muscles and some firmly lodged between bones (Corbett, 1957). In another dead adult tiger the liver and lungs were pierced at many places by porcupine quills (Burl’er, 1955)*. Evidently tigers in the Trans-Caucasus, Middle Asia, and southern Kazakhstan also suffer injuries from this rodent. The diseases suffered by tiger in the wild have not been studied. In zoological gardens they suffer from all infectious diseases characteristic of cats as well as those of other species of carnivorous animals. In one menagerie a Ussurian tiger suffered from food poisoning and died of it on the thirteenth day. The infection was transmitted by consumption of the flesh of a diseased horse (Vyshelesskii, 1948). Tigers also die of pasteurellosis, paratyphoid, carnivore distemper, etc. Two tigers caught in southwestern Tadzhikistan harbored five to seven tapeworms (Taenia bubesei) in the small and large intestines; these worms had formerly been recovered from African lion (Chernyshev, 1953; in the past the ranges of tiger and lion often overlapped). Toxocara mystax has also been detected in this predator (Mozgovoi, 1953). Toxocariasis is often noticed among tigers in zoological gardens. In an Amur tiger the following four species of helminths were found: Paragonismus westermanni, Physaloptera praeputiale, Dirofilaria ursi, and Uiteinarta sp. (У. Abramov, 1962). From a female caught in Tadzhikistan on June 26, 339 ticks belonging to four species were collected: Rhipicephalus turanicus (greater part), Boophilus *Not in Literature Cited — Sci. Ed. 149 187 calcaratus (three), Hyalomma detritum (seven), and Н. marginata (Chernyshev, 1958). Primor’e tigers are [pele eet by the tick Dermacentor silvarum (Pomerantsev, 1946). The wolf is a competitor of tiger everywhere. It is known that at places in the Primor’e where tigers live, wolves are absent or very few in number; it is thought that tigers destroy wolves (Kaplanov, 1948; G.F. Bromlei). From the diligence with which a tiger chases dogs, one may presume that it will hunt wolf just as tenaciously. Leopard could also be a competitor of tiger; usually these two cats do not live in proximity to each other. Sometimes bear and yellow-throated marten trail behind a tiger to scavenge carcasses; jackal, jungle cat, and hyaena similarly follow tigers in Middle Asia. In the Far East, in the Amur valley, a male wapiti killed by a tiger was found by a Fig. 83. Track of a three-year-old tiger (male) in fine snow. Sankhobe River basin Sikhote-Alin preserve. February, 1960. Photograph by G.M. Veinger. 2 149 188 brown bear who buried it in the snow (K.G. Abramov). In the Khabarovsk region in years of food scarcity (pine nuts and acorns) bear have no chance to fatten and do not remain long in their lair. Instead they set out to hunt more often than usual for wild boar and moose, thus competing with tiger. Under such circumstances bears sometimes chase young tigers off their prey (V.P. Sysoev). Crows and ravens invariably gather around the remains of a tiger kill. Small predators are not afraid of a dead tiger; in 1950 in Tigrovaya Balka preserve (Tadzhikistan) a tiger carcass was found which had been gnawed at by jackals. Population dynamics. \t was mentioned earlier that the tiger population in the last century decreased everywhere and at present it has disappeared altogether in many parts of its range (Uzbekistan, Kazakhstan) or is only found as an intruder (Trans-Caucasus, Turkmenia, Tadzhikistan, southeastern Trans-Baikal, and Amur district). The reason for the rapid decline in the population of tiger was persecution by man and especially the indirect effects of his intervention (felling and burning of forests including the tugais, drainage, cultivation, and burning of reed floodplains, reduction in population of wild ungulates, and other activities). The main reason for the death of the last of the tigers in Kazakhstan was the extensive burning of reeds in the lower reaches of rivers during April and May every year, which destroyed all vegetation there in the 1930’s through the 1940’s (Sludskii, 1950). Tigers disappeared from the lower reaches of the Amu-Darya mainly because of fires (Pokrovskii, 1951). In Tadzhikistan their population decreased as a result of rapid cultivation of river floodplains (for example, Vakhsh). When the agriculture of Middle Asia and southern Kazakhstan suffered great losses as a result of wars (most of the large irrigated fields and cattle were destroyed) in the thirteenth and eighteenth centuries, the tiger population rose significantly. A similar phenomenon has been observed in China and India. Following major social upheavals (wars, revolts, and revolutions), which disturbed the economy of these countries, the tiger population began to rise. For example, in China the number of tigers rose significantly in the years of the National Liberation War which ended in 1949. Along with a gradual reduction in number of predators, temporary changes have also been detected in their population due to migrations, and mortality during unfavorable conditions, mainly scarcity or difficulty of access to food. In Sikhote-Alin in years of abundant yield of pine nuts and acorns, the wild boar population increases, and, subsequently, the number of tiger. The latter migrate there from northeastern China (Manchuria) and the Korean Peninsula. In the Amur district after the large-scale mortality of wild boar in 1955, tiger disappearance was almost complete (G. Modin). 150 Fig. 84. Tiger tracks on wet coastal sand. Ta-Chingou area, Sudzukhin preserve. May, 1965. Photograph by V.E. Prisyazhnyuk. In the lower Amu-Darya a perceptible reduction in tiger numbers took place after an epidemic plague broke out among wild boar in 1938, deci- mating 50% of their population. After the emigration of wild boar from Amu-Darya delta in 1945, tigers disappeared from there almost completely. In some years, a perceptible rise in tiger population has been noted in the tugais on the right bank of the Pyandzh. Such phenomena occurred in 1936 and 1950. In the latter year local hunters killed four tigers there in three or four months. The observed increase in tiger population on the Pyandzh’s right bank was associated with fires in the tugais on the left bank (in 150 Afghanistan; Chernyshev, 1958). The tiger population may drop after winters 190 Fig. 85. Track of a tigress on shoulder of а forest road. Upper Tudo-Vaku River in the Iman basin. Sikhote-Alin. September, 1967. Photograph by E.N. Matyushkin. of abundant snow and snow crust prolonged into early spring. It is known that in such winters adult tigers suffer intense hunger and even die of emaciation (see above), but the young suffer even more since the mother cannot provide adequate food. Tiger mortality may possibly result from epizootic diseases breaking out among feline and other carnivorous animals. The main factors influencing tiger populations, apart from man and his activities, are climatic conditions which affect the population of ungulates serving as their prey, ease of access to food, and possibly epizootic diseases. Field characteristics. When moving fast a tiger appears to be a mono- chromatic light brown. It usually paces, striding almost step by step (Figs. 82, 83, 84, and 85). The stride is 50 cm to 80 cm long or even 100 cm in particularly large males. In galloping, jumps measure 4.0 to 5.0 m in length and sometimes more. If the track had a length® of 20 cm or more, “’The length of a track is measured from the front point of the impression of the longest digit to the rear point of the sole, i.e., the rear of the padded portion (hunter’s terminology). Invariably the size of the print of a front paw is given, since the forepaws are larger than the hind ones. 151 191 the animal is large; that of a male is usually 16 cm x 14 cm and of a female 15 cm x 11 or 12 cm. Depending on the width of the “‘heel’’ impression and other characteristics, four categories of adult and sex groups can be roughly identified: 1) ‘“heel’’ width 10 to 15 cm—adult male weighing 200 kg or more; 2) “‘heel’’ width 9 to 12 cm—adult tigress weighing 150 kg or more; 3) ‘‘heel’’ width 6 to 8 and 9 cm—cub up to three years of age weighing 70 to 120 kg, in rare cases up to 150 kg; and 4) “‘heel’’ width 3.0 to 5.0 cm—cub weighing 30 to 60 kg (V.K. Abramov, 1961). The tracks of a tigress are not so rounded as those of a male and the digital impressions are thinner (Fig. 87). The thick middle digits are particularly prominent in the male. Moreover, the sex of the animal can be readily determined from urine marks. The female urinates on the ground, Fig. 86. Track of a tigress with cub. Old cut-over at head-waters of the Ulakha, Sikhote-Alin. December, 1969. Photograph by A.G. Pankrat’ev. 152 Meee Yow Bes Fig. 87. Schematic prints of front and left hind paws of a tigress; measurements 149 mm x 154 mm and 131 mm and 150 mm. Ta-Chindzhan mountain. Sudzukhin preserve. Spring of 1947. Sketch by G.F. Bromlei. as a result of which a thawed patch is visible in the snow, while the male sprays urine on any prominent object such as logs, mounds, etc. Young males also leave a fairly prominent print but its depression is not great because of their lesser weight. An adult tiger’s pace is fully equal to a man’s stride while that of a young cub is shorter. When a three-year-old cub follows its mother, the tracks almost overlap; when mother and cub move independently two sets of prints are distinguishable. The presence of younger cubs is evident from multiple prints in a small area because they are playful. The well-being of adult animals can also be judged from the sharpness of paw imprints. In the case of well-fed tiger the pug marks are sharp and deep; contrarily, in an emaciated one they are flat and shallow and wrinkles in the pads discernible (Fig. 87). Numerous fecal deposits are a sure sign of imminent approach to a site where the predator has consumed a prey. Feces are black or brown in color and undigested remains of the prey distinguishable—fur, bone, etc.; they are generally cylindrical, about 4.0 cm in diameter, and slightly pointed at the rear end. But when meat and blood predominate in the food, the feces are semiliquid, shapeless, and blackish. Both types of feces exude a sharp, putrid, long-lasting odor. That the tiger has fed in this area is confirmed by the discovery within a short distance of scattered vertebral bones, long bones and sometimes even the skull of a large ungulate. The roar of tigers is heard rarely even at places where they are quite common. Only during rut is the animal’s roar common. Usually the first roar is protracted, repeated in quick succession, and terminates in three or 193 153 Fig. 88. Remains of three-year-old tiger cub killed and eaten by a brown bear. Shukhi-Pokto preserve, Khabarovsk region. 1956. Sketch by N.N. Kondakov from a photograph by Krivopusk. four short roars. Sometimes a low throaty ‘‘ah-oun’’ or ‘“‘ee-oh-unnu’’ is emitted. When enraged the animal issues a hollow snarl and produces coughlike sounds with an intense exhalation of air through an open mouth (Baikov, 1925). Moreover, tigers sometimes produce a high pitched sound which could be described as ‘‘muuk’’ or ‘‘puuk’’ and is somewhat similar to the bugle of a wapiti in rut. Some hunters and even zoologists assume that tigers entice wapiti through such a sound. In India at places where there are many sambars tigers imitate their call to attract them into ambush (this opinion requires confirmation). A contented and resting tiger rumbles* a long time but, on being frightened, issues a characteristic ‘‘vuuf’’. Tigers generally attack their prey silently. (A.S.) Practical Significance When tigers were numerous they caused some damage to animal husbandry. For example, in 1905 a pair of tigers living on the Karatal River (southern Pri-Balkhash) killed about 30 camels and 100 horses (Shnitnikov, 1936). As late as 1929, in the lower reaches of the Ili, tigers destroyed over 200 *Myrlychet in Russian original; probably a misprint for murlychet— Sci. Ed. 154 194 head of cattle (М. Sh., 1930). In 1938 a large tiger in Parkhar (Tadzhikistan) destroyed in one month 23 large and small cattle on a collective farm. Cattle particularly suffer from this predator in wintering sites located among reeds by rivers. Tigers inhabiting the Chu River attack wintering cattle, mainly cows, and more rarely camels (Gern, 1891). With the arrival of Russians in Middle Asia, 1.е., from the middle of the nineteenth century, intense hunting of tigers began. Moreover, their destruction was encouraged. In the 1870’s a prize of 25 rubles was awarded for every tiger killed and at the beginning of the present century raised to 50 rubles. Even as late as 1929 in Semirech’e a special decree was issued for killing tigers. Tigers could be killed by any method and a bounty of 100 rubles collected. All these measures and some other factors led to the rapid extinction of the predator. In the 1930’s and later in Middle Asia tigers continued to attack cattle from time to time only in Tadzhikistan and the Amu-Darya delta, but this loss was extremely small. Formerly, tigers sometimes harmed cattle even in the Far East. In June, 1869 at Troitsk, on the shore of Lake Khanka, tigers killed in one month 32 cows and horses (Przheval’skii, 1870). In the last decade the damage caused by tigers to cattle has been negligible even in these regions. At present the damage done to game husbandry by tigers is negligible because of their small population. Moreover, zoologists who have studied tigers in the Far East in the last decade consider them useful to game husbandry, since wolves are absent or few in number at places where tigers are present. Tigers do kill a few ungulates but, unlike wolves do not drive them away (Salmin, 1940; Kaplanov, 1948; G.F. Bromlei). Several instances of attacks on man by tigers have been described т. the Soviet Union in the last century. They occurred mainly in Middle Asia (excluding Turkmenia), Kazakhstan, and the Far East. Usually animals shot and wounded or chased by hunters attacked, and only very rarely did an attack occur without provocation. Nevertheless, in the lower reaches of the Syr-Darya a tiger reportedly attacked and tore to pieces a woman collecting firewood, and an unarmed military officer passing in June through reed thickets (“‘Vernenskii Grazhdanin,’’ 1880). Instances of tiger attacks on shepherds have been reported from the lower reaches of the Ш (A.M. Nikol’skii, 1885). Attacks on man in the Far East were recorded in the middle and third quarter of the last century. In 1867 on the Tsymukha River, tigers mutilated and killed 21 men and wounded 6 others (Przheval’skii, 1870). In the present century no tiger attacks on man without provocation have been reported (Kaplanov, 1948; Abramov, 1960; G.F. Bromlei). In preserves where tigers are conserved today they pose no threat to man because they are not hunted here and wild ungulates are bountiful. True man-eaters, i.e., animals for which man is a regular, if not the only Nn Fig. 89. Young tiger pressed to the ground with a ‘‘fork’’. Photograph by V.G. Toropov. prey, are now absent in the Soviet Union. This is due to their relatively small population and the abundant availability of ungulates at places where they live. In India, however, animals regularly attacking man were numerous in the past and some still exist today. In the middle of the last century man- eaters were a common phenomenon in some regions of central India. In the Monula region alone they killed 200 to 300 humans in one year and decimated entire villages (Jerdon, 1874). Even in 1924 tigers killed 603 humans in British India (K-li, 1926). One tigress in Nepal killed 200 humans and in Kumaon over a four-year period before she was herself destroyed —234 (Anderson, 1964). In the central regions of India about 155 196 50 men were killed by tigers every year in the last decade (Berten, 1954*). Man-eaters are usually animals compelled to feed on humans, an un- natural food for them, being unable to hunt healthy, strong animals. The reason for such behavior in nine out of ten cases is a wound, and in the remaining case old age (Corbett, 1957). The wound might be the result of an unsuccessful shot by a hunter or the predator’s attack on a porcupine. In China attacks on woodcutters and coachmen have been reported in Kirin province (Baikov, 1925). Instances of tigers entering cabins and carrying off adults and children are also known. Once a tiger attacked and killed three farmers in a forest but did not feed on the carcasses. In the southern part of China tigers killed 60 men in a short period of time (Allen, 1938). Attacks on humans by tigers have occurred in recent years in Yunnan province, even as late as 1949 (A.A. Sludskii). Tigers are valuable fur-yielding animals but the importance of their skin as an item of clothing is negligible today. Throughout the Soviet Union, before hunting was banned, only ten animals were caught annually and their skins usually retained by the people; three or four were killed in Tadzhikistan . and five to eight in the Far East. Only one to four skins were received for curing (G.F. Bromlei). Formerly, most skins were retained by the hunters or bought by collectors. Only three tiger skins (from Kirgizia steppe) passed through the custom houses at Petropavlovsk, Presnogor’ kovsk, Om’sk, and Korkovsk from 1857 to 1861 even though dozens of tigers were killed annually in the nineteenth century (Krasovskii, 1868). The number of tiger skins coming from the USSR in relation to the world output is almost negligible. The world output from 1807 through 1910 was estimated as 500 skins (Kaplin, Ivanov, and Patushenko, 1955); Russian fairs annually received only 30 to 50 skins up to 1914. In the 1920’s around 60 tigers were caught in the Soviet Union, including two or three live ones (Solov’ev, 1926). In these same years the annual catch in Amur and Primor’e provinces was 25, in Korea 25, and in northeastern China 50 to 60 (Baikov, 1925). Amur tiger skins are especially prized in the world market. The rising price of tiger skins is one of the factors responsible for their persecution. For example, in 1760 a large tiger skin in Orenburg fetched 13 rubles; a medium-sized one 8 rubles, and a small one 5 rubles (Rychkov, 1762); in the early 1900’s the price exceeded 100 rubles (Zarudnyi, 1915) and even rose to 200 rubles in Middle Asia. The skins of Far East tigers were even more expensive. In the Primor’e region a tiger skin cost 70 rubles in 1886 and as much as 100 to 200 rubles in 1895 (Silant’ev, 1898); at the same time sable skins sold for 5 to 25 rubles each. In 1927 and 1928 in the Far East a tiger skin was valued at 300 to 500 Mexican dollars (Baikov, *Not in Literature Cited— Sci. Ed. 156 197 1928). The present market price of a cured tiger skin varies from 4 to 35 rubles depending on its quality. The carcass of a tiger is no less valuable than the skin; even in the 1930’s tiger carcasses were exported to China, where they were sold to pharmacists. The people of China, Semirech’e, and the Kul’dzhinsk region eagerly bought the heart and bones of tiger (Gern, 1891). In 1929 and 1930 in the Far East tiger carcasses fetched 3 rubles 50 kopecks per kg, and the bones 15 rubles per kg. During this same period in northeastern China, 10 Mexican dollars were paid for 16 kg of carcass (Baikov, 1928). A hunter could thus receive up to 1,000 rubles for a single adult tiger. Formerly in the Far East, caught animals were not skinned but were frozen whole and dispatched to the nearest town where they were bought by Chinese marketeers and transported to interior China, mainly Peking and Tientsin. Chinese medical practitioners highly prized the carcass of an adult male and its various parts. Medicines were prepared from each organ which cured not only different diseases, but also cowardice, emaciation, etc., and talismans were made from them. In the middle of the last century Kazakhs generally cut out the claws of a tiger and stitched them on wearing apparel for children; claws were considered charms which drove evil spirits away from a child (Karelin, 1847). Even in 1958 tiger bones were sold in China. A single animal yielded sometimes up to 30 kg of bones. Particularly prized were the bones of the leg, especially the patellas (Sludskii, 1966). Various body parts of the tiger have been used in Korean medicine. Both Chinese and Koreans enjoy tiger meat. From very early times tigers have been caught live and exhibited in circuses or kept in menageries. Roman emperors often displayed their tigers. In the Middle Ages tigers were trained in the east for hunting big game (Simashko, 1851). From 1260 to 1280 Kublai Khan, grandson of Genghis Khan, regularly used tigers on hunts in China (Marco Polo, 1955). In ancient times in the Trans-Caucasus, Middle Asia, and Kazakhstan, tigers were hunted for sport and for the purpose of their destruction. In Azerbaidzhan, Armenia, and Georgia, czars and aristocracy used to arrange hunting parties for tigers. Tiger hunting was very prevalent during the period of Mongol rule. Mongolian aristocracy and military detachments hunted tigers in order to perfect military techniques and to inculcate bravery in the soldiers. In the Middle Ages Mongolians and peoples elsewhere regarded the tiger as a superior being and were afraid of it (Grekov and Yakubovskii, 1950). With the decline of the Great Khans, hunting of tigers also declined, but with the arrival of Russians in Middle Asia in the nineteenth century, it was revived оп a large scale. Soldiers from border regions, special ‘‘hunting units’’ of the army, and individual hunters, mainly army officers, began 198 hunting tigers. Tracking and catching tigers (and wild boars) helped instill bravery in the Kazakhs and soldiers. Regular hunting of wild boars and tigers on the Syr-Darya greatly influenced the qualities of courage of local garrisons, and inculcated valor and military dexterity in the soldiers who ‘“became fearless because accustomed to facing danger, and later such garrisons distinguished themselves in many notable battles in the Syr- Darya region’’ (‘‘Vernenskii Grazhdanin’’, 1880). The Kazakhs of Syr-Darya and Amu-Darya garrisons hunted tigers by lying in wait with their guns or by shooting animals beaten out of rushes. The hunt was sometimes conducted on horseback or sometimes small groups charged as in a bayonet attack. Almost every such attack resulted in the death or wounding of soldiers and officers. The local people of Middle Asia hunted tigers by attaching a knife to a “‘spring”’ on a track or at the site of a killed calf. A flexible pole with the knife tied to it was lowered to the ground and fixed with a simka®! in such a way that the animal on touching the latter released the pole. As the pole straightened the knife ripped the animal’s underbelly. In the Amu- Darya a large automatic type of trap used by northern trappers was employed. In the tugai a strap tug supported a tiny pole fitted with sharply pointed knives. A careless tiger on touching the strap released the tug and the pole with the knives hit the flanks of the animal. Kazakhs from time to time set up log mats with nails and knives on the outer logs. All these methods were usually ineffective and the animal often escaped with wounds; nevertheless one or two were caught in a season (A.I., 1888). More commonly tigers were caught in large portable traps and shot. The most prevalent method of hunting tigers was to set up around the remains of its kill one or several guns which went off when the animal touched a string. As in the previously described methods of hunting, the powerful animal, though shot, was rarely killed on the spot. Injured, it fled and later became dangerous to man. In the last century Kazakhs sometimes caught tigers by poisoning the remains of its prey with a poison prepared from the roots of Eminium lehmanni. Sometimes hunters lay in wait for the animal at the site of its kill or on its path after constructing a wooden platform protected from all sides. The platform was fitted on poles firmly driven into the ground. The tiger was shot from the platform and if it attacked, killed from above with a bayonet or large knife. Sometimes Kazakhs joined three such platforms together and firmly tied sticks on top. A thin strap was fastened to the top of this cage. Several men sat inside the cage equipped with guns and other sharp weapons. Another method was to go to the animal’s den wearing А fine string used in making various traps. 199 157 the protective device described by Gern (1891). Kazakhs also hunted tigers 158 in groups of 30 to 40 men mounted on horses, killing the beast with bow and arrows (Eversmann, 1850). In the Far East local hunters (Udekheits, Evenks, Orochets, and others) did not hunt tigers, as they regarded the species with reverence and caught one only under exceptional circumstances (see below). In the nineteenth century Russians bagged tigers with both bow and gun. A bait such as a slaughtered wild boar or roe deer was placed in the animal’s path. More often a dog served as bait and loaded guns were set up around it (Kulagin, 1923). The animal, wounded by the automatic guns, sometimes attacked the men chasing it. More rarely, the animal was poisoned with strychnine. Tiger skins were used in making carpets and rugs. In ancient Rome live tigers were imported from Persia and Armenia. In the eighteenth century animals from Bukhara were brought into the menageries of Russian czars (Р.Г. Rychkov, 1762). There were still tiger catchers in the Ussuri territory from 1902 to 1906. Some farmers caught live tigers with their bare hands, using neither cage nor trap, and tied the animals up with ropes (V.K. Arsen’ev, 1949). There was a particularly great demand for live Ussurian tigers in the 1920’s. At that time an adult animal fetched 1,000 pounds sterling (about 10,000 gold rubles) and a cub of up to one year, 100 pounds. The demand for live tigers within the Soviet Union and abroad continues even now. A young Ussurian tiger in the 1950’s to the 1960’s fetched up to 500 rubles or more. On the international market a Ussurian tiger in the mid-1960’s cost about $4,000 and a southern Chinese tiger about $3,000. The great demand for live tigers resulted at one time in the capture of almost every cub produced annually. Usually a group of four or five hunters on finding the tracks of a tigress with cubs pursue them, often for several days. On closing the gap the hunters yell and chase the animals, shooting in the air in order to separate the tigress from the cubs. This chase sometimes covers 5 to 10 km. The weaker of the cubs begins to lag behind its mother, who continues to move forward. Having lost sight of its mother, the frightened cub strays from her path, turning sideways. Dogs, kept leashed until that moment, are then set on its course. ' The released dogs frighten the cub until it seeks shelter under an upturned or wind-felled tree, or in rocks, etc. On reaching the cub and restraining the dogs, the hunters roll and pin the cub to the ground with poles with forks at the end. They then tie its legs with straps or rope loops and muzzle it with a pre-prepared muzzle (K.G. Abramov, 1956). This is acommon method of catching tigers live, but quite often another is employed. Sometimes the cornered animal is poked by one or two men 157 200 Fig. 90. Tied cubs. Primor’e region. Winter of 1952/1953. Photograph by М. Nazarov. with the butt of a rifle or a padded jacket or bag. Before it can spring for an attack other hunters overpower it and, holding it by the nape of the neck and paws, pin it to the ground where it is trussed up (Shklyar, 1935; Reutov, 1957): Having caught the first cub of a litter, one of the hunters stands guard over it while the rest start chasing the remainder until the entire litter is caught. Rarely is a cub capable of eluding the hunters. A group of coastal: tiger catchers detected 17 litters comprising 48 cubs from 1936 through 1957 and caught 45 of them (V. Abramov, 1962). The efficiency of the methods described above for catching tigers can be judged from the following data. From 1930 through 1940 on the Iman and its tributaries alone, 40 cubs and even 9 adult tigers were caught (Sysoev, 1955a). One tiger hunter caught 21 and killed 18 (up to 1935); another caught 36 in his lifetime, and still another caught 16 and killed 8. Catchers from Loumo from 1947 through 1955 caught 24 tigers of which some weighed 116 kg (cubs are caught live up to the age of three years inclusive, when they quite often weigh over 100 kg). In the remote past, among peoples of the Far East, and also China, Vietnam, Japan, Mongolia, Korea, India, Nepal, Indonesia and other countries, there existed a special tiger cult. Tigers and dragons were depicted 159 158 201 as objects of art in palaces of rulers, huts of cultivators, cabins of animal catchers, and on altars as idols. In the Altai, in excavations of the Pazyryk burial mounds, circa 500 to 300 B.C., some lifelike drawings of tigers were found together with animals with the body of a tiger or the head of a tiger on the body of an eagle or vulture (Rudenko, 1953). In the palace of Toprak Kal, in ancient Khorezm, tiger illustrations have been found (Tolstov, 1951). In the forests of the Far East animal catchers and those seeking ginseng raised idols in order to propitiate the tiger. Among the Chinese, Udekheits, and other peoples, tigers aroused a special mystic fear and reverence; the dark pattern on the head of a tiger resembles the hieroglyph ‘‘уап-да,’ which means ‘great emperor’’ (Fig. 91). According to popular belief, the spirit of a ruler lived in the body of a tiger as a means of expiating his sins. Formerly, Udekheits killed a tiger only if it attacked their camps. The tiger cult (mafa cult) among these peoples resembled the cult of the ‘‘master’’ of nature—of the taiga and the universe. Myths of a totemistic nature were associated with the tiger (Ivanov, Levin et al., 1956). The tiger cult prevailed even among the Orochets. They killed a tiger in accordance with the laws of their native land, only to avenge the killing of their kinsmen. Nanaiets also had a special reverence for the tiger. The members of one family traced their ancestry to this animal. All the foregoing beliefs reveal the important role tigers formerly played in the life of peoples of eastern Asia. As a gift of nature and history the tiger is of paramount scientific importance, which is increasing every year due to the animal’s rapid decline. The tiger is no longer dangerous nor harmful to man in the Soviet Union. Every available measure will have to be implemented to prevent its total destruction. Such measures include a total ban on tiger hunting and severe restrictions on catching of cubs throughout Soviet territory. In the Khabarovsk and Primor’e regions all types of tiger catching were totally banned from Fig. 91. Hieroglyph `‘уап-Ча’’ (right) meaning ‘‘great emperor’’ and dark-colored pattern resembling it on head of a tiger. Sketch by N.N. Kondakov. 202 1947 through 1954. In 1954 a temporary relaxation was granted for catching cubs. In the Primor’e region in 1956 shooting and catching of tiger were banned throughout the year and throughout the territory for a period of five years. At the same time a penalty of 5,000 rubles (old currency) for unauthorized tiger killing was imposed. Hunting of tigers had been banned even earlier (1952) in this region. The ban on catching and killing of tigers in the Khabarovsk and Primor’e regions continues to the present day (1968). However, beginning in 1960 special licences have been issued for catching a limited number of tiger litters. Measures for protecting the Far East tiger have produced positive results. The decline in its population, counted at the beginning of the 1940’s, has been arrested at about 120 to 140 individuals (see above). In Tadzhikistan the killing of tiger year-round was prohibited in 1957 but, sad to say, the ban came rather late. Tiger conservation requires an international agreement with governments of neighboring countries in which this animal still survives (Korean Democratic Republic, South Korea, China, Afghanistan, and Iran). It is necessary to set up large sanctuaries in the regions adjoining the Korean Democratic Republic, Chinese People’s Republic, Afghanistan, and Iran, and to enlarge the Sikhote-Alin, Sudzukhin, and Suputin preserves (the upper Suputinka, originating in the Dadyan’-Shan range in the western spurs of Sikhote-Alin). A preserve is necessary in Birobidzhan for preserving tigers in the Shukhi-Pokto range, and game refuges should be set up in other places where tigers still survive and steps taken toward the conservation of ungulates in them, which serve as food for the predator. Raising tigers in zoological gardens would help to increase their number in the Soviet Union. The implementation of these and other measures would ensure the survival of this most interesting animal in our fauna; if unheeded, this cat will be totally extinct in a few years. Tigers are protected in several foreign countries. In the Chinese People’s Republic the hunting of Amur tigers inhabiting the northeastern province is totally banned. Measures for the protection of tigers have also been effected in the Korean Democratic Republic. In India tigers are protected in preserves and in a series of long-term game refuges. In the Union of Burma tigers are likewise conserved in long-term game refuges. The Javan tiger* in Indonesia is protected in a sanctuary in the western part of Java. (A.S.) *Now extinct—Sci. Ed. ~ 160 203 BARS* [LEOPARD] Panthera pardus Linnaeus, 1758 1758. Felis pardus. Linnaeus. Syst. Nat., 10th ed., p. 41. Egypt. 1856. Felis tulliana. Valenciennes. С. К. Acad. Sciences, vol. 42, р. 1039. Ninfy, 40 km east of Smyrna (Izmir), Asia Minor. 1857. Felis orientalis. Schlegel. Handl. d. Dierkunde [sic], vol. 1, p. 23. Korean Peninsula. 1862. Leopardus japonensis. Gray. Proc. Zool. Soc. London, р. 262. Northern China (ascribed in original description to Japan where leopards are absent and were never present). 1867. Felis fontanieri. Milne-Edwards. Ann. Sc. Nat. Zool., vol. 8, p. 375. Near Peking. 1867. Leopardus chinensis. Gray. Proc. Zool. Sec. London. Mountains west of Peking. Non Felis chinensis Gray, 1837. 1878. Felis leopardus. Sclater. Proc. Zool. Soc. London, р. 289. Persia (?). ©The correct name in the Russian language is ‘‘bars’’ and not leopard. Intense efforts were made in the last decade to introduce the latter name into Russian literature. The word ‘Багз’ has been in popular usage and literature for several centuries (at least from the sixteenth century) and right from the beginning it referred to the Caucasian leopard, i.e., to P. pardus. Among Russian peoples this name is used throughout the Caucasus. The word “*panther”’ was used for the first time at the end of the nineteeth century and only in the territory of the Caucasian preserve among hunters; the word ‘‘panther’’ was introduced during the period of the great imperial hunts from Petersburg. Russian peoples called, and continue to call the animal ‘‘bars’* throughout Turkmenia, in the Soviet Far East, and in northeastern China (Manchuria). This name was used in Russian zoological classics of the eighteenth, nineteenth, and early twentieth centuries (5.1. Ognev, for example), and is still used by many well- known contemporary writers. It has long been listed in Russian dictionaries with a reference to P. pardus (**. . . like a desert bars, ferocious and wild ...’’ or ‘‘. . . the desert’s permanent guest—mighty bars ...’’ Lermontov; or ‘‘wearing a bars skin,”’ etc.). In the distant past, in the eighteenth century, as can be seen from Pallas (1811), but evidently even before, the word “‘bars’’ was popularly used for irbis [ounce] (Uncia uncia) in Siberia—and later in Semirech’e, and Middle Asia. This is quite natural considering the similarity of these two species and such confusion is not uncommon. Furthermore, in the fur trade of the seventeenth century, references also occur to the skins of *‘irbizy’’. In the recent period (nineteeth to the early twentieth century) in Russian zoological literature the name ‘‘snow leopard’’ (identical in concept to the English, German, and French names) and “‘irbis’”’ (ounce) (from Turkey and Mongolia) has been firmly established for Uncia uncia. The term ‘’Багз,” as earlier, is thus retained for P. pardus. There is no sense in changing the name ‘‘irbis’’ to ‘‘bars’’ nor introducing the new name “‘leopard”’ for **bars’’; neither is there any scientific justification for doing so. Neither is there zoological or philological justification, nor any practical convenience achieved. Such changes would only lead to misunderstandings, confuse nonspecialists altogether, and needlessly clutter up the language. (V.H.) 204 1903. Felis villosa. Bonhote. Ann. Mag. Nat. Hist., vol. 11, р. 475. Amur Gulf. 1904. Felis pardus grayi. Trouessart. Cat. Mammalium, р. 268. New name for chinensis Gray. 1914. Leopardus pardus ciscaucasicus. Satunin. Consp. Mamm. Imp. Ross., vol. 1, p. 159. Opredelitel’mlekop. Ross. Imp., vol. 1, p. 159. ‘‘Forested region of northwestern Caucasus (montane zone of the Kuban district.’’) 1927. Panthera pardus saxicolor. Pocock. Ann. Mag. Nat. Hist., ser. 9, vol. 1, p. 213. Astrabad, Iran. 1959. Panthera pardus dathei. Zukowsky. Zool. Garten, vol. 24, p. 348. Sandabad, southeast of Shiraz, southern Iran. 1964. Рапйега pardus transcaucasica. Zukowsky. Zool. Garten, 28, 4, 158. ‘‘From Georgia to Armenia and Azerbaidzhan’’ (sic!—V.H.). (V.H.) Diagnosis Color yellow and reddish-yellow with small black rosettes and spots with a well-defined light colored center. Body length less than 160 cm, condylobasal length of skull less than 250 mm, and length of upper carnassial tooth less than 29 but more than 21 mm. (У.Н.) Description Size large. Much smaller in size than the tiger or lion, the leopard, snow leopard (ounce), and cheetah are nonetheless bigger than the remaining Russian cats and represent the next largest species in the family (Fig. 92). Body long, with rather short iegs; very agile, light and slender, with a long tail (length more than one-half length of body), and a relatively small rounded head. Forehead bulges, facial portion moderately long, and ears short, rounded, and wide-set. Eyes small, with round pupils. Mane or longer hair on dorsal side of neck and cheek absent; hair of very uniform length throughout the body, relatively short and close-fitting, and not luxuriant even in winter. Type of hair frequently responsible for slender appearance of animal. Body, however, very strong and muscular with powerful and broad front paws. ‘*Pelage short, coarse, and sparse. There are about 3,000 hairs рег cm? on the back, with only four underfur hairs to one guard hair. A characteristic difference is apparent in the length and thickness of black (in the spots) and yellow hair; the former (as in all fur-bearing animals) is thinner and longer. Yellow guard hairs and 30 mm long and 121 microns thick but the black hairs 40 mm long and 96 microns thick. The length of yellow top “лолешоу ‘МУ Аа зицитеа “| права vsayjuvg ‘рледоэр ueisvoned ‘т эеа и 161 Fig. 92. Far East leopard, Panthera pardus orientalis Schleg. Vil’nyus Zoological Garden. Photograph by B.B. Marm. hair (four categories) is 26, 25, 24, and 22 mm, and thickness 109, 92, 56 and 37 microns; the corresponding measurements for black hairs are 30, 31, 29, 27 mm, and 89, 64, 57, and 49 microns; the length of yellow underfur is 20 mm and thickness 21 microns and of black underfur 24 mm and 32 microns’’ (B.F. Tserevitinov; specimen from Middle Asia). Differences between the summer and winter coats vary in different subspecies but generally small. In Amur leopards the length of hair on the back in summer: is 20 to 25 mm and in winter 50 mm. The color of leopard is distinguished by a general light background of yellow or reddish-yellow color with black spots of two types—solid or annular (rosettes). In the center of the latter a light-colored field occurs, corresponding fairly well in color to the main background of the coat. The black ring may be complete but is usually interrupted at two to five places, 1.е., it consists of two to five separate spots grouped into one ring. Transverse (vertical) stripes are not present but sometimes individual spots on the back fuse into short longitudinal stripes. The outlines of the spots are well defined. The general background color varies individually, geographically, and 206 seasonally. Among Soviet leopards it changes from a fairly dull light yellow or yellowish-gray to a bright yellowish-red with a golden tinge, but may also be light gray or almost white or whitish. The color is most vivid on the back, and often on the dorsal part of the neck and head. It gradually lightens and loses the yellow and red tinges on the flanks, and is white on the abdomen, inner side of the limbs, the throat, below the neck, extremity of the legs, the underside, and the terminal half and upper side of the tail. The color contrast between the back and rest of the body is variable and depends on the general intensity of coloration. This contrast is less sharp in light-colored forms than in reddish-colored animals. The extent and character of spottiness is variable. Spots are usually pure black in color but may also be brownish. The total number and density of disposition of spots is likewise variable. Also variable are the size and form of solid spots and size and structure of annular spots, i.e., the number of their constituent elements, their rounded or oval form, relative numbers in the general pattern of skin and disposition on the body, and color of the central field (corresponding to the yellow background of the skin or darker), etc. The anterior part of the muzzle is devoid of spots and small dark markings occur only in the region of the vibrissae and a dark spot on the lower lip in the corner of the mouth. On the cheeks, forehead, between the eyes and ears, and along the top and sides of the neck, solid black spots of various sizes, but generally small, are present; on the dorsal side of the neck the spots are somewhat elongated. Back of the ears black. Annular spots occur on the back and flanks. Spots on the spine are usually elongated or form a pattern of elongated-annular and large elongated solid spots. Sometimes individual spots fuse into a short longitudinal stripe. Annular spots, consisting usually of a few sections and measuring up to 50 to 65 mm in diameter, are distributed throughout the flanks, on the top of the shoulders and on the thighs or in their upper parts. While the annular spots on the flanks descend to the abdomen and usually comprise the predominant type of spots (some are distributed lower), their distribution on the shoulders and thighs is more variable. Thus on the shoulder and forelegs they may be present together with solid spots and even descend to the forearm or even be absent on shoulder. They may also cover the entire region of the thighs or only their upper part. Solid spots occur all over the body, wherever annular spots are absent, but sometimes are present together with several or many annular spots. They are few on the flanks, close to the spine, and more numerous toward the abdomen. The legs and abdomen are covered with solid spots. The spots on the outer side of the legs are large toward the top and smaller down below, becoming tiny freckles on the wrist and paw. The top of the tail and part of its underside are covered with large annular 163 207 or solid spots. The vibrissae consist of black, white, and half-black, half- white resilient hair up to 110 mm long. The claws are a light waxy color, highly compressed from the sides, steeply curved, and very sharp; their length along the outer arc reaches up to 55 mm (front paw). Claws on the hind paws are smaller and not so sharp. Seasonal color variability and the nature of the coat in Soviet leopards are distinctly manifested but not to the same degree in all forms. In spite of long hair, up to 50 mm in winter, the pattern of spots is sharply maintained as in summer. The main background color in winter is paler and duller than in summer, but quite vivid even in winter in brightly colored forms. In pale forms it may even be whitish, resembling the color of the ounce. Individual color variability is generally confined to changes in vividness of color of the main background and the pattern of spots (see above). Two strikingly different color types—duller or more vivid—are seen in some populations. These are not connected, or almost not connected, by inter- mediates (Flerov, 1935; Tsukovskii, 1959; also see ‘‘Geographic Variation’’).8 Color macromutations affecting spottiness, described from India (Pocock, 1941), are not known among Soviet leopards. Melanistic animals (‘black panther’’) are quite common in some parts of southern Asia (Sikkim and Nepal—P. p. pernigra Hodg., and Java—P. p. melas О. Cuv.), but absent or extremely rare within the Soviet Union.*4 Young leopards have a very light background color, which is yellowish-white on the back and whitish on the flanks. Their spots are brown or brownish- chestnut, similar to the color of the back of the ear. Spottiness is very dense, however, and hence the animal appears quite dark. Geographic color variation pertains to shade of color and vividness of the general background (from a very light grayish-white or pale yellow to a bright and full reddish-chestnut) and in several details of the nature of the spot pattern (see ‘“Geographic Variation’). The skull of a leopard differs from that of a tiger in many features (see description of tiger), primarily size. Even in very old animals the skull exhibits less prominent “‘predatory’’ features and is more ‘‘catlike’’ (see characteristics of the family). In leopard the interorbital and postorbital constrictions are smaller, the cranium larger, zygomatic arches less separated, facial part relatively less developed, etc. 8*Грезе, however, do not represent a dual-phase color but extreme types of variations. The absence or rarity of animals with intermediate colors is explained by the nonavailability of a large series of skins in museums. “One report (Flerov, 1935) is available pertaining to southern Tadzhikistan, but quite possibly it could also pertain to the snow leopard. The absence of melanistic animals in the Middle Asian and Caucasian parts of the geographic range, which are known for their aridity, particularly the former, is quite expected. 208 The skull of leopards is characterized by the following features. Оп the whole it is relatively massive, fairly long with rather close-set zygomatic arches, and quite low. Upper line of profile from highest point in interorbital region to rear runs relatively gently, but forms a very steep arc toward the front. Frontal area almost flat or with faint longitudinal depression. Crests, especially sagittal and lambdoidal, highly developed (Fig. 93). Rostral width (above canines) in adult animals wider than postorbital constriction and more or less equal to interorbital width. Zygomatic arches strong and zygomatic process of temporal reaches base of postorbital process of jugal. Postorbital process of jugal does not reach level of lacrimal foramen. Lower anterior region of orbits slightly thickened and bluntly rounded toward the top. Infraorbital foramen fairly large, its maximum diameter exceeds width of bony part between it and edge of orbit. Nasals long and fairly uniformly narrowed in posterior part. Hard palate relatively narrow and extended; its width much shorter (by 35 to 50%) than its length. Posterior edge of bony palate in the middle of interpterygoid vacuity (along the suture between the palatines) with small pointed projection. Tympanic bulla (see Fig. 23) relatively large, greatly inflated, with a fairly smooth surface. Anterior chamber (p. ectotympanica) small and occupies much less than one-half of volume of entire bulla. Its boundary (usually poorly visible) is shifted in an anterolateral direction. Anterior margin of bulla lags far behind glenoid fossa and postglenoid process, by not less than one-half width of interpterygoid vacuity. Distance between inner walls of tympanic bullae more than width of interpterygoid vacuity. Jugular foramen large. Paroccipital process massive and closely adheres throughout to posterior wall of tympanic bulla and does not forma free falcate process. Canines relatively thin at base but long and sharp. Additional fifth tubercle absent on anterior outer part of upper carnassial tooth. Sexual dimorphism reflected in small overall size and slender skull structure of female. In males, as a result of the far more intense development of masseter muscles, the entire cranium is more compressed. Frontal part of cranium and postorbital constriction (“‘waist’’) long and highly compressed; cross section (“‘postorbital width’’) roughly equal to inter- orbital width. This isthmus in females is usually short and the compression there less: width of this region more than interorbital width. Crests, especially sagittal and lambdoidal, less developed and teeth on the whole somewhat weak; canines short and thin, diameter of alveoli small, and alveoli not so deep in females (Pocock, 1941; for Indian animals).® ®5Sex related differences in the skull may be significant. In the absence of adequate and correctly dated material, erroneous taxonomic conclusions have sometimes been drawn. д У Эк И! >>”, < КА b ke т ee № № Any я Al DN QO , д а SN: SSS S Я ow МЕХ = N SS Г SS Г \ = о = GAN aa tS orig gp TE aly iy LU, 5 Y/ РР 2 Z Los ИТ Г ЗМ. АМ ARW\ oA } = NN \\ 164 Fig. 93. Skull of leopard, Panthera pardus ciscaucasica Sat. No. S 51945, Zoological Museum, Moscow University. Kopet-Dag, 1935. Sketch by N.N. Kondakov. 210 Age-related changes in the skull of leopards are the same as those in tigers. In young animals the skull has a relatively short and broadly expanded cranium, small facial part, close-set zygomatic arches, and rather smooth interorbital and particularly postorbital constrictions. Crests are either absent or slightly developed. The skull lengthens with age. Age-related changes in the skull of leopards differ markedly from those in tigers in that their range is smaller, 1.е., the morphological contrast between the early and last stages of postnatal development is jess. This is explained by the slightly more “‘catlike’’ structure of the leopard skuil and the elimination of the later stages of refinement seen in the tiger. Leopards exhibit the beginning of this process, which progressively develops in various species of the family and ultimately leads to a decrease in contrast between the early and late age-related morphological stages. Geographic changes in skull structure are little known but evidently involve changes in overall measurements. Data on size of leopards inhabiting the Soviet Union®° are very few and do not cover the entire range of variation in these features. Body length (male and female) varies from 120 to 171 cm, tail length 75 to 102 cm, length of hind foot 24 to 26 cm (four animals), length of ear about 7.5 to 8.0 cm, and height at shoulders 50 to 78 cm. Adult animals weigh 32 to 60 Ке.87 Weight may possibly reach 75 kg (Stroganov, 1962). References to 117 and even 130 kg (Baikov, 1927 and 1929), often quoted in literature (Ognev, 1935), are positively erroneous (see below). This is underscored by the fact that they are said to pertain to the rather smali Far East form (specimen from northeastern China), although representing a record for that country, yet fali within the range of variation of linear measurements of the species (body length 145 and 151 ст).88 A ‘‘very obese’’ male killed near Sud- zukhin preserve on August 12, 1945, although measuring 128 cm in length, weighed only 44 kg (G.F. Bromlei). The known maximum for Amur leopard within the Soviet Union is 48 kg. Two Amur leopards, males of “‘average dimensions’’ and ‘‘moderate obesity,’’ killed in February and March, 1927 ®According to Dinnik, 1914; Satunin, 1915; Bil’kevich, 1924; Baikov, 1927; Ognev, 1935; Stroganov, 1962; material furnished by G.F. Bromlei, N.K. Vereshchagin, A.G. Pankrat’ev, and A.A. Sludskii; and collections of the Zoclogical Museum, Moscow University and the Zoological Museum, Academy of Sciences. 8’/These measurements, taken ‘‘in the flesh’’, cover all the published data for Russian animals and those obtained from skins and mounted specimens except one, 1.е., 183 cm for a Caucasian leopard (skin from Lenkoran; Satunin, 1915). *8Data furnished by М.А. Baikov on the size and weight of leopard from northeastern China are extremely contradictory. For example, apart from the data given above, a weight of up to 160 kg (**10 poods’’) has been quoted; 96 kg (‘6 poods’’; 1915) is more common. The length of these animals generally reaches 155 cm and the tail length not more than 86 cm, but a figure of 95 cm has also been adduced for the same region (1927). 166 AA near Vladivostok, weighed 32.2 and 32.8 kg (Ognev, 1935). The weight of a large female evidently from northeastern China was 42.5 kg; two other animals (femaie and male) weighed 25 and 29 kg (Chou, 1958). A leopard from Kopet-Dag weighed 60 kg; the forms there are evidently very large.°° The following data provide some idea of how Russian leopards compare with Indian ones in size (P. p. fusca—whole of India except Kashmir, Nepai, Sikkira, Baluchistan, and Sri Lanka). Body iength of four large males from India 137, 130, 128, and 128 cm, and taii length 84, 84, 76, and 92 cm (Pocock, 1941). Average measurements of 11 males from Sri Lanka: body length 126.5 cm, tail length 85.9 cm, height at shoulders 64.3 cm, and weight about 56.34 kg (124.1 English pounds). The largest male from Sri Lanka (an exceptionally large animal, largest of ail animals known) and a body length of 142 cm, tail length 96.5 cm, and weighed about 77.2 kg. Average measurements of seven females from Sri Lanka: body length 104.7 cm, tail length 77.5 cm, and weight 29 kg. The largest female had a body tength of 114 cm, tail length 84 cm (total length 198 cm), and weighed 44 kg (Phillips, 1935). Maximum skuil iengih of five adult males from India 232 to 250 mm and condylobasai length 207 to 221 mm; corresponding measurements in six females 177 to 201 mm and 159 to 185 mm (Pocock, 1941). Greatest skull length in males 193 to 256 mm and in temaies 180 to 218 mm; condylobasal length in males 186 to 223 mm and in females 170 to 188 mm; zygomatic width in males 123 to 172 mm and in females 116 to 135 mm; muzzle width above canines in maies 53 to 65 mm and in females 47 to 53 mm; length of upper tooth row in males 65 to 79 mm and in females 60 to 68 mm.” (V.H.) Systematic Position While lion and tiger are very closely related species, the leopard is notably distinct from them. From the viewpoint of systematics, the leopard is closely related to jaguar (P. onca). This view is supported by the coat pattern and craniological features. Of these two species, leopard represents a more specialized form (Haltenorth, 1937), 1.е., a more progressive one. However, it is possible that leopard is closer to tiger than is jaguar, which thus -compietes the specialization series in the genus (lion—tiger—leopard — jaguar; Tsarapkin, 1939; see also characteristics of the genus and ‘“Systematic 8%*A large male up to 7 poods’’ (112 kg) from this region (Bil’kevich, 1924) is an outright misstatement. **According to the data available in literature, material of the Zoological Museum, Moscow Univeisity and the Zoological Museum, Academy of Sciences (М.К. Vereshchagin), and data furnished by А.О. Pankrat’ev (Vladivostok) and М.1. ishadov (Ashkhabad). 212 Position’’ of tiger). In any case the skull of leopard, compared to that of tiger and lion, does not exhibit many “‘primitive predatory’’ features; instead it reflects a significant step toward the evolution of a distinct cat type of skull. (V.H.) Geographic Distribution Leopards are found in forests, parts of forest steppes (savannas), and montane regions of Africa and southern Asia, the Near East and the southern half of eastern Asia. Geographic Range in the Soviet Union The range in the Soviet Union represents the northern rim of distribution of the species and consists of three isolated sections —Caucasian, Middle Asian, and Far Eastern, which are connected to the south outside USSR boundaries. The Middle Asian section, in turn, is broken into a few individual pockets (Fig. 94). In the Caucasus the range (reconstructed) covers essentially the whole of the country except steppe areas (Fig. 95). Leopards used to occur in mountains and foothills covered by forests and also in desert hills devoid of forests. In the west the range commenced at Tuapse, possible even slightly more westward, but probably did not reach Novorossiisk. Intersecting at this meridian the western offshoots of the Great Caucasus, the range boundary encompassed the northern foothills and ran east and southeast up to the extremity of the mountain range—toward Buinaksk and Makhachkala. In the foothills leopards were confined to forests and at places where forests merged into the plain or descended to very low levels. Thus, in the western Caucasus leopards were known from Samursk station, Elisavetpol’sk village and ccrtain other sites to the south and southwest of Maikop; in the central Caucasus in the region of Ordzhonikidze (Vladicaucasus); in the east, at Evdokimovy on the Argun, near Makhachkala town itself (Agach-aul), and at Karabudakhkent and Gubden, southeast of Makhachkala.?! *!L_eopard skins sold in Kizlyar in the eighteenth century (Pallas, 1811) were most probably brought from the hills but quite possible leopards also intruded from the foothill forests into Terek valley forests, which were formerly far more extensive than at present, closely associated with the foothills and hill forests, and abounded in ungulates. In general, however, there is no information relating to the distant past (up to the nineteenth century) for the Caucasus; yet indubitably leopards were present everywhere in the northern Caucasus wherever there were forests. It is highly probable that in the remote past (before the appearance of Russians), the animal was generally far more extensively distributed in the Caucasus than has been recorded. Partly, as at present, these occurrences were in the form of intrusions along the periphery or into the plains of the eastern Trans-Caucasus, but it is impossible to distinguish such past instances from the fairly normal permanent habitation. о © 2 y 16 5 0 3 } <] ail а al [Pp < [] suit 1 лы, Lake ealkhash iy Wale eae Te | ee 1000km 750 250 500 Fig. 94. Reconstructed range of leopard, Panthera pardus L. in the Soviet Union (scale in km). Dots depict some sites of distant intrusions (for more details see Figs. 95, 96, and 97). V.G. Heptner. 168 214 Along the Great Caucasus leopards occurred up to the upper forest boundary and even higher. On the southern slope they descended to the foothilis and even to the plains as in the north. In the eastern part of the mountain range the animals were known in Alazan valley at Lagodekh, Zakatal, and Nukhu, and recorded in the west at Adler, Sochi, Tuapse, and Bzybi. They therefore descended in this region to the sea. п the Trans-Caucasus the range of leopards covered all of the montane region of Little Caucasus, including in the west the Trans-Caucasian expanse between Rioni and Kur’, and the Pontic range in the southernmost part of the coast (Satunin, 1915).°? There is no direct evidence about the | | | | ee =< aE ii il UN _ ЩИ , | | | п | ND il o Fig. 95. Range of leopard, Panthera pardus ciscaucasica Sat. in the Caucasus (scale in km). 1 — probable boundary of reconstructed range in the historic period; 2— region of fairly regular, at places very rare, occurrence of leopards by 1950; 3— intrusions of leopards into the steppe; 4—some sites where a few wandering animals were caught from the 1930's to the 1950’s outside the territories shown by 2; 5 — possible occurrence at Terek in the eighteenth century and before. V.G. Heptner. *Pontic range refers here to the northernmost extremity of this mountain range to the south of Batumi, then a part of Russia. 168 169 215 occurrence of leopards in the marshy lowlands and the forest expanses of Kolkhidsk lowland, but there is no doubt that they prowled there, though rarely, because of scarce game. The distribution also covers the Talysh and Lenkoran lowlands. In the steppe part of the eastern Trans-Caucasus leopards were evidently absent. However, wandering animals were seen even in forestless steppes and semidesert plains far away from the mountains. A case of transgression into Sal’ yan was recorded at the end of the nineteenth century (Radde, 1899) and one in 1946 into the densely settled part of the Apsheron Peninsula (Bil’gya, 25 km east of Baku; Dzhafarov, 1946); some transgressions into the “‘central Mugana reeds’’ have also been recorded (Vereshchagin, 1959). By the 1950’s to the 196075 the range of leopards in the Caucasus had shrunk greatly, the population of the animal became negligible, and actually on the brink of total extinction. This was due to the ruthless killing of the animal, which had never been particularly abundant. The rapid and steady disappearance of leopards, especially in the Great Caucasus, has mainly occurred over the last 80 to 100 years, particularly in recent decades, since even at the end of the nineteenth century it was still sighted throughout most of the Caucasus. Not even the establishment of the Caucasian reserve in the 1920’s could restore the dwindling population of this animal in the western Caucasus. From the 1930’s to the 1950’s leopards, though extremely rare, were still sighted in the forest regions of the western (Kuban) Caucasus, mainly in the Caucasian preserve and adjoining areas. In the 1930’s in this region there were two pockets of leopard habitation—one in the upper reaches of the Kuban, Kishi, and Beloe (Atamazha, Tybga, Dzhemaruk, Chugush, and Assara ranges), and the second in the upper reaches of the Sochi, Khosty, Golovinka, Bzyby, and Shakha, which flow into the Black Sea. It is possible that a few leopards inhabited the montane regions of the eastern Caucasus above Zakatal and Lagodekh and possibly the heavily forested sections of Dagestan associated with them. In the rest of the Great Caucasus, on both northern and southern slopes, over the last decade only a few stray intrusions (once in several years or decades) have been reported, sometimes at places in the western, central, and eastern parts of the range, far from the places specifically mentioned above (Fig. 95)*.33 *Misnumbered **96°’ in Russian text—Sci. Ed. Occurrences at Zelenchuk and Arkhyz are known. Distant intrusions include Makhach- kala and the Apsheron Peninsula (see above). One leopard was killed at Lars on the Georgian Military Highway in 1900 (Beme, 1941) and another roughly at the same place in the 1940s. In February, 1949 a pair of leopards was seen at Galaskha in the Groznyi region. Other instances of distant intrusions are known, for example, towards Tskhinvala (southem Ossetia, 1952) (see also Vereshchagin, 1959). 216 In the Trans-Caucasus (not including the Great Caucasus range), in the 1950’s and early 1960’s leopards still inhabited the Talysh, Karabakh, and Zangezur ranges in the north to the Kel’badzhar region and Murovdag, and westward along the mountains to the north of the Araks valley through Nakhichevan Autonomous Soviet Socialist Republic almost to Yerevan (to the Saraibulag range and the Veda region). They also occurred in Kubatlinsk and Zangelansk regions of Azerbaidzhan and possibly from time to time in the northern forest regions of Armenia (north and northwest of Lake Sevan). This section of leopard habitat, at least in its southern portion, was associated with the geographic range in Iran and intrusions southward occurred from there. The probability of such intrusions has diminished in recent decades. In the mid-1960’s leopards were already absent in Armenia, except probably for some individual animals in the Zangezur range. On the Talysh, however, leopards were resident throughout the 1960’s and evidently not mere intruders. Almost every year one or two leopards are killed there to the west of Astar and Lenkoran (road to Lerik). Two animals were shot in October, 1969 west of Astar (V.S. Lobachev). On the whole, by the middle and end of the 1960's leopards had already practically disappeared or were passing through their last days in the Trans- Caucasus and the Little Caucasus. They still occur, though very rarely, at places where there are several tributaries from Iran, 1.е., on the Zangezur (southeastern Armenia and southwestern Azerbaidzhan) and persist on the Talysh. Some animals may have also intruded from these areas into other parts of the Trans-Caucasus. Leopards do not enter the Soviet Union from Turkey. In the Great Caucasus some leopards are evidently still met with along the southern slope of the eastern section (Azerbaidzhan and eastern Georgia) and some strays probably live in the Kuban region of the Caucasus and on the southern slope of the western half of this range. Leopards are absent in the rest of the Great Caucasus and in certain areas have not been seen for quite some time.”4 The Middle Asian habitat of leopards (Fig. 96) in the Soviet Union is associated with the distribution of the animal in the Near East, in Iran and Afghanistan, and represents the northern extremity of its range. It is divided into several sections. In the west. in Turkmenia, a large section covers all of the Kopet-Dag from its western and northwestern (Kyuren’-Dag) rim to the extreme east, i.e., roughly to Artyk railway station (about 59°30’ Е. *4The range in the Caucasus, in addition to works cited in the text, is based on Dinnik, 1914; Satunin, 1915; Heptner and Formozov, 1941; Nasimovich, 1941; Vereshchagin, 1947 and 1949; Burchak-Abramovich and Dzhafarov, 1949; Dal’, 1954; Kotov and Ryabov, 1963; Alekperov, 1966; and other sources; as well as original data compiled by V.G. Heptner. See section **Biology’’ for more details about the Caucasus and the reduction of the range there. 170 217 long.), where the mountain extends beyond the Soviet Union. An isolated very small section is formed by the Little Balkhan upland (774 m above sea level) and a slightly larger one by the Great Balkhan (1,880 m above sea level). Both of these are isolated from each other and from the Kyuren’- Dag section by desert plains —the so-called Balkhan passes (‘‘corridors’’), each 25 km wide. Obviously, the leopard population in these sections is likewise isolated. Yet, in spite of the Turkmenian leopard’s affinity for mountains (it does not enter the plains for more than a few kilometers), one could nevertheless assume that formerly, when leopards (like mountain sheep) were widespread in Kyuren’-Dag and their population high there, some exchange of animals must have occurred between Kopet’-Dag and the Balkhans (Fig. 96). Farther to the east there is a separate section (within the Soviet Union) of leopard habitat to the west of Badkhyz in the Gyaz’-Gyadyk upland (up to 1,000 m) extending from Iran. There the animal reaches in the north, to Pul’-i-Katum on the Tedzhen and is found in the middle of Badkhyz in the Lake-Er-oilan-daz depression (ur. Namak-saar) and in the Kyzyl-Dzhar gorge. Farther east, again after some interruption, leopards occur in the mountains east of the Kushka River, between Kushka and Kashan and the upper Murgab. Toward the north there they are not found beyond the mouth of the Kushka on the Murgab and probably do not even intrude in that region. Information about the habitat of leopards along the Tedzhen and Murgab outside the sites discussed above is erroneous and mainly due to mistaking cheetah for it. North of the Great Balkhan leopards are not encountered and occasional information about their occurrence there also pertains to cheetah. In the Kopet-Dag leopards occurred (or still did even in the 1940’s) right up to the foothills of the range and sometimes even emerge onto the premontane plain. Thus, intruding from Iran, they stayed from time to time in the premontane plain slightly east of Artyk, even at Mean and Chaach (south of Tedzhen city) and moved toward the Caspian Sea in the Atrek basin. Similarly, in southern Turkmenia leopards from Gyaz’-Gyadyk went east into the Er-oilan-duz depression (data for Turkmenia from V.G. Heptner). A separate section of the Middle Asian range of leopards is situated on the right bank of the upper Amu-Darya and Pyandzh. There leopards live in low (up to 1,100 to 1,400 m above sea level) mountains in the west up to Babatag (between Surkhan-Darya and Kafirnigan), in the north to the Gissar valley (upper reaches of the Kafirnigan), and in the east to the foothills of the Darvazsk range and the Parkhar region on the Pyandzh, where their range adjoins that of the ounce [snow leopard]. Within the boundaries described above, leopards have been sighted in Bahatag, Aktau, Oktau, Karatau, Terglitau, the Bal’dzhuan mountains, in the regions of Koktash, 1эщ4эН “OA 'С8азаея U19]S9M JY] UO) вАзеа-пигу рие церчеАд до Yue 348 и VO ио!3э1— / ($14 YoINSusyD) лэлг qudinyy Joddn рие vyysny usamjaq зитдипош — 9 ‘zkyypeg иг uotssaidap znp-uejio-1q рие sureyunow yApedn-,zeAg — © (©) Зв -.иэтаАМ чим 3eq-jadoy — р ‘чецея aI] — < ‘иецу ея 12210 —Т ‘(WY ит э1е9$) BISY э1ррид Ul еб 92150пр5'> зправа влащию] ‘рледоз| JO эВиел рэзэпц$иоээ$ ‘96 ‘ЗЫ ЕЕ: WHOS OGL } о + 219 Shaartuz, Dangarin, Mikoyanabad, and Parkhar (Flerov, 1935; Chernyshev, 1950; Ishunin, 1961). All information about the residence and catch of leopards in Semirech’e (Shnitnikov, 1936; and others) is erroneous and refers to ounce. Even random intrusions of leopards into this part of Middle Asia should be regarded as exceptional. 145 ql ПВ rT h : | 80 0 80 150 240Кт oo a oe пежеиа п Fig. 97. Range о leopard, Panthera pardus orientalis Schleg. in the Far East (scale in km). A—northern boundary of fairly permanent habitation in the 1950’s to the early 1960’s (data of С.Р. Bromlei; N.V. Rakov, 1965); B—boundary of region of maximum permanent habitation in 1950 (С.Е. Bromlei); С— intrusions in the middle of the last century (1—Tyrma River, and 2— Bureinskii mountains); D— some intrusions in the 1910’s to the early 1920’s (3 — иррег reaches of Iman, and 4— Sitsa); E—some intrusions from 1939 to 1962 (5-30 km south of Vyazem, 6— left Pokkhorenok, and 7—Bikin midreaches). Section of the course of the Amur between ‘‘а’” and ‘‘b’’ represents region of leopard intrusions in the middle of the last century (Maak, 1859). For intrusions of leopards into Trans-Baikal, see Fig. 94. V.G. Heptner. 171 220 In Middle Asia the reconstructed range of leopards described above, unlike the situation obtaining in the Caucasus, mostly still corresponds to the range at the end of the 1960’s. Leopards, however, are absent in the northwestern part of the Kopet-Dag (in Kyuren’-Dag) and became extinct, or almost so, in the Great Balkhan by the early 1950’s. The population throughout the range also underwent radical changes (see below). The Far East range of leopards (Fig. 97) within the Soviet Union is contiguous with its range in northeastern China and the Korean Peninsula. The predator has been found in the Ussuri territory and in parts of the Pri- Amur and Trans-Baikal. In the middle of the present century (1940’s to 1960’s) the region of fairly permanent habitation, including regular intrusions (see below) was limited to the far south of Ussuri region. The northern boundary of the range there commences on the coast of the Sea of Japan and Dzhigit Bay at 44° N. lat. and runs south at a distance of 15 to 30 km from the coast to the latitude of Valentine Bay (43°10' М. lat.). There it turns steeply westward, north of the Suchan basin, then north to encompass the source of the Ussuri (Daubikhe River) and its right bank tributary, the Ulakhe, part of the basin of the right tributary of the Ulakhe Noto, and the Shetukhe, the right tributary in the upper reaches of the Ussuri (G.F. Bromlei, Rakov, 1965). Here the boundary turns westward toward the bank of [Lake] Khanka at the level of the state boundary (Figs. 97 and 98). South and west of the above line leopards occur everywhere, at least, they did in the past, in the Khanka lowland (south of the lake). The asso- ciation of Ussuri leopards with mountains is fairly definite but evidently not as rigid as in the case of Turkmenian leopards. In the area described leopards are rare, especially in the more northern parts; they are confined more to places where wild sika deer live or where deer husbandry is practiced. Thus in the 1950’s leopards were observed at Razdol’nii (50 km north of Vladivostok), Gamov Cape, Kita Bay, ‘““Kedrovaya Pad’ ’’ preserve, Tuman Bay, Glazkov, along the rivers Steklyanukha, Maikhe, and Lefa, and elsewhere (G.F. Bromlei; for more details, see also Rakov, 1965). The unique feature of leopard habitat in this part of the Soviet Union is that this area is one of occurrence of solitary wandering animals, groups in pairs, or a mother with cubs. Definite data on breeding of leopards throughout this region are not available and it is possible that the animal does not reproduce in this part of the Soviet Union except perhaps in ‘*Kedrovaya Pad’ ’’ preserve (southwest of Vladivostok). In general, it consists of animals originating from the western* part of China. This does not contradict the fact that here they are a settled species and relatively *Sic: should read ‘‘eastern’’—Sci. Ed. 7 Э Lo Sovetskaya Gavan’ x 4 [i= 4 Fig. 98. Present distribution of leopard, Panthera pardus orientalis Schleg., in Ussuri territory. 1—boundary of region of permanent residence by 1971; 2—best sites of habitation; 3—region of very rare and temporary intrusions beyond zone of permanent occurrence; 4—directions of individual intrusions from China during 1930 to 1970. (S.P. Kucherenko.) common in places. Such a position is generally quite normal for the periphery of the range of distribution. This explains, probably, some ambiguities in establishing the boundaries of the leopard’s range in the southern Ussuri region. These, however, pertain to frequencies of sightings. A comparison of range data for the past and present brings us to the conclusion that the region of permanent habitation of leopards in the Ussuri territory has not altered significantly over the last hundred years. With such an insignificant region of more or less regular habitat leopards cover a large area with long intrusions which, at one time, contributed to an incorrect depiction of the range of the species in the Soviet Far East and in Trans-Baikal. This is mostly explained by the fact that older zoologists 222 usually relied on inconsistent information from local peopie and were partly influenced by the erroneous statements of Pallas (1811) about ounce between the Uda and Amur.® Thus it was even thought that leopards inhabited the Amur region to the same extent as tigers, 1.е., the entire course of the Amur, the shores of the Okhotsk and Tatarskii seas, and Sakhalin Island (Schrenk, 1862). At present there is justification to consider all the areas where leopards have been found outside the region described above as regions representing deep and rare intrusions. Furthermore, ali these areas represent intrusions not from the south of the Ussuri territory but from northeastern China. Only the intrusion into the Sitsa River region may be an exception. The following are the northernmost known poinis of leopard occurrences: southeastern Trans-Baikal (very rare); Kuchugai on the lower Gazimur, a tributary of Argun in eastern Trans-Baikal (Cherkasov, 1887; Baikov, 1927); Nerchinsk- Zavodsk region of Chitinsk district (1952; A.A. Sludskii); Bureinskiy mountains (at the Amur; Radde, 1862); Tyrma River, a tributary of the Bureya (evidently in the upper reaches in these same mountains; Middendorff, 1867); and along the Amur between the mouths of the Sungari and Gorin.”° All these represent very distant and, evidently, rare intrusions in the past century, in part in the middle or the first of it. In the 1910's and 1920’s intrusions of individual animals were observed in Sitsa in the Terneya region (coast of the Sea of Japan in the proximity of 45° М. lat.) and in the upper reaches of the Iman (around 46° М. lat.; Arsen’ev, 1926; Baikov, 1927). In the middle of the present century intrusions are known in the middie course of the Bikin, the Khor basin (the Left Podkhorenok), on one of the right tributaries of the lower reaches of the Ussuri (30 km south of Vyazem), to the mouth of the Bir and on its middle course (Rakov, 1965), and others [sic]. In the past, intrusions were evidently more frequent and extended farther, but even at present leopards move quite far, even in Trans-Baikal (additional data on intrusions and distribution in the range are given under ‘‘Biology’’). Old information about the habitat of leopards in Sakhaiin (Schrenk, 1858; Nikol’skii, 1889) is undoubtedly erroneous. Similarly, assumptions about leopard habitat in the eleventh and twelfth centuries at the mouth of the Don and in the southern Russian steppes or forest-steppes, based on an interpretation of the “‘fierce animal’’ of old Russia as leopard (Pidoplichko, Older researchers (Maak, Schrenk, Przheval’skii, and Nikol’skii), following Pallas, usually applied the name Felis irbis to the Far East leopard. This has led to confusion even among some present-day zoologists (Zhitkov, 1934) and is a purely nomenclatural error. Ounce are absent in the Far East and were never present there; all data pertain to the leopard. °°In the expanse *‘from the mouth of the Sungari ending slightly above the mouth of the Gorin, the Tuzemtsy [local people] were aware, apart from tiger, of still another big species of the cat genus’’ (Maak, 1859). 174 223 1951; Mavrodin, 1964; and others), are unfounded (See section on tiger and also Heptner, 1969). The discovery of parts of the lower jaw of a leopard, used as an amulet, in the ruins of one of the Greek Black Sea colonies cannot serve as adequate proof (Vinokur and Khotyun, 1967). This should merely be considered an import from Asia Minor, with which Black Sea Greeks conducted an active trade. On the whole the range of leopards in the Soviet Union is in the process of undergoing reduction, at places very rapidly. The population is partly supplemented by animals migrating from adjacent territories, where they are more commen and suffer less persecution (Iran, Afghanistan, and China). If measures are not implemented soon for the conservation of leopard, this species will totally vanish from the Soviet Union in the near future. Geographic Range outside the Soviet Union The range outside the Soviet Union includes northeastern China (Manchuria), the Korean Peninsula, eastern China (evidently not Kansu), the eastern border of Tibet,?’ Indochina, Malacca, Java, Kangean Isiands,?* Burma, India, the Himalayas, Pakistan, Afghanistan, Iran, Asia Minor, the Arabian Peninsula, Sinai, Egypt, Morocco, Algiers, Tunisia, and the whole of Africa south of the Sahara (Fig. 99). At present the range everywhere is drastically reduced. In the Quaternary period leopards lived in France, Italy, England, Germany, Belgium, Spain, Portugal, Switzerland, Yugoslavia, Hungary, and Rumania. Some discoveries point to the occurrence of this species in Europe up to and including the Neolithic period (Pidoplichko, 1951). Within the USSR leopard remains have been found in the Middle Quaternary formations of the Tashik-tash caves in the Baisunsk mountains, which represent the very fringe of the present-day range. In strata of the fifth to the second century B.C. a lower jaw was found at Olvia (Nikolaevsk district). This was evidently the remains of an imported animal. (V.H.) Geographic Variation Geographic variation in leopards is well-marked, which is natural considering "In general, usual references to *‘Tibet’’ are erroneous. Leopards are evidently not seen in the uplands and in the ranges surrounding them from the north. Residence in eastern Tibet (Kam and others) has been established with ceztainty. п Sumatra leopards are absent and all data about their residence there should be considered erroneous. At the same time the break in the range at Malacca-Java appears very strange and probably leopards were present in Sumatra in oiden times. The existence there of a very small number of animals has been assumed even by recent authors (Chasen, 1940). References to the occurrence of leopards in Japan (Dal’, 1954) are unsubstantiated. 224 е i SSS SSeS A SS ew es EY 7 Sa eee и, Saas cee 1 ss SSeS SO OOo ea oe se Ge an ШЕгоопатыя ЕБЕЕЕСЕВ [oeemge eo coal [= ae ees or TT | eA ( | м li ть о ЕЕ: == 0 1000 2000 3000 4000 Кт Fig. 99. Reconstructed range of leopard, Panthera pardus Satunin (scale in km). In the northeast the boundary also includes region of intrusions. Information about Sumatra appears dubious. V.G. Heptner. the vastness of their range and the diversity of natural conditions within it. Nevertheless this variability is not as sharp as might be expected under these conditions, although extreme forms differ rather notably. The main background color of the coat may be very pale—pale yellow or even grayish-white—on one side of the range, and bright—red or reddish- chestnut on the other. A transition between these extremes evidently-does not occur sharply anywhere, although in some parts of the range a subspecies or a different category lives in relative proximity. 175 In other features variability is reflected in intensity of the black tone of the spots, their number, size, density of disposition, relative number of rosettes and their position on the body, the color of the center of these spots, etc. For some forms a dual-color type has been noted. In some populations 225 of the humid regions of southern Asia the percentage of melanistic forms (‘‘black panthers’’) is high (Sikkim and Nepal —P. р. pernigra and Java— P. p. melas), while in other predominantly arid zones melanism is totally absent or an extremely rare phenomenon. On the whole the geographic variability of leopards distinctly conforms to Gloger’s rule. This pertains not only to the general shade of color, but also to the total area of black patterning and number of melanistic forms. Within the USSR and adjacent sectors of Asia, Bergmann’s rule is not manifested so distinctly, but in the range as a whole much smaller forms are associated with warmer and more arid regions. At the same time, for example in East Africa, these animals are very large, judging from the skull size. Such big animals have not been reported in the Soviet Union to date (Best et al., 1962). Geographic variation in leopards has not been studied adequately. At present a large number of subspecies have been described and the number of distinct and even accepted forms far exceeds those actually existing. Almost all the subspecies described are based on color characteristics; however, as pointed out before, coloration exhibits extreme individual variability. Individuals exhibiting extreme color variability from a given region have sometimes been placed in separate subspecies (see later). Geographic variation for the entire species has not been revised in accordance with our present level of knowledge. At the same time contravening modern concepts (also in nomenclature), the practice continues of maintaining as distinct old forms and even separat- ing new ones on the basis of casual data and even stray photographs and skin descriptions. Disagreements and changes in the concepts of geographic variation of the species are consequently great indeed. Hence 15 subspecies were recognized from Africa in the 1930’s (over 25 names; G. Allen, 1939); of these, many pertain to the eastern part of the mainland and three to south Africa (Roberts, 1951). Later (Ellerman, Morrison-Scott and Hayman, 1953), all South African animals were placed in one nominal form (Egypt), 1.е., the African expanse from north to south is now regarded as populated by one single subspecies. This positive critical approach contradicts the tendency of extreme subdivision. A large number of leopard subspecies are recognized from Asia (12; Ellerman and Morrison-Scott, 1951 and 1966). In the Soviet Union various zoologists have generally designated five subspecies. In reality only two exist. 1. Amur leopard, Р. р. orientalis Schlegel, 1857 (syn. villosa, japonensis, fantanieri [51с; fontanieri]|, chinensis, grayt) Size not large. Coat fairly soft, with long (on the back 30 to 50 mm and on the abdomen 70 mm) and dense hair. Main general color type bright and lustrous. Winter 176 226 coat varies from fairly light yellow to dense yellowish-red with a golden tinge or rusty-reddish-yellow. Color on flanks and outer sides of legs lighter. Spots pure black color; light-colored centers of circle of spots (‘‘rosettes’’) somewhat darker than main background color of skin. Spots numerous, 1.е., spottiness prominent (Fig. 100). Summer pelage shorter and brighter with more vivid coloration pattern. Skull small, with narrow interorbital region (on average, width about 20% of condylobasal length); postorbital constriction distinct, short, and in the form of an isthmus; nasal pointed at posterior ends and zygomatic arches relatively massive. Measurements of male (six): body length 107 to 136 cm (M 125); tail length 82 to 90 cm (M 86); length of hind foot 24 to 27 cm (M 25); and height at shoulders 64 to 78 cm (M 72). Maximum length of skull 204 to 232 mm (M 213); condylobasal length 186 to 200 mm (M 189); zygomatic width 129 to 144 mm (M 138); interorbital width 34.3 to 39.9 mm (M 37.8); т width 36.8 to 45.0 mm (M 40.2); and length of upper tooth row 67.8 to 68.7 mm (M 68.0) (Stroganov, 1962). Weight of males of moderate size 32 kg (Ognev, 1935) and of large ones 48 kg. This weight may even reach 60 to 75 kg.” This subspecies is encountered in the Amur and Ussuri regions. Outside the Soviet Union Amur leopards are found in northeastern China, probably in the south to Peking, and the Korean Peninsula. Amur leopards are clearly distinguishable from leopards inhabiting Turkestan and the. Caucasus. The differences between Amur and Chinese leopards are complex and not fully understood. Thus some false notions have arisen. As mentioned above, the range of color variability in Ussuri leopards is quite broad and differences in color intensity and brightness of extreme forms fairly sharp. Differences in vividness of coloration between the winter and summer coats are also distinct. This sometimes creates the impression of the existence in the Soviet Far East of two forms of leopard (subspecies), differing in color. п northeastern China (Manchuria) out of 18 animals killed and 18 skins the following are the measurements of the two largest (Baikov, 1927 and 1929): body length 145 and 151 cm, tail length 95 and 82 cm, and maximum length of skull 221 and 255 mm. The weight of these two animals has been shown as 117 and 130 kg which, of course, appears erroneous (see “*Description”’ above); this is the weight of adult Amur tigresses. (The author perhaps meant pounds and wrote kilograms by mistake—? On converting Russian pounds, the weight would be 46.8 and 52 kg and English pounds 53.0 and 58.9 kg, which in general accords with other data.) The following are the dimensions of three adult males from northeastern China: body length 106, 110, and 106 cm; tail length 76, 70, and 70 cm; length of hind foot 23, 22. and 24.3 cm; height of ears in all three animals 7.5 cm; and weight 25, 42, and 29 kg respectively (Shaw, 1958). ‘roujday “OA Aq ч4ел3о3оца `Аизлэлии Г] MODSOY] “WNasN] [е910[007, ay] JO (0190 1013 suauIIoads [PY 1809 JOWIUINS ‘UOITaI 105$] ‘16189 $ ‘ON 909 JaIUIM “pogy ‘aoutAosd Зипушен ‘eaoy илэЧиоц ‘00166 $ “ON “EZ6T “I9IUIM “VaIOY илэчиои Jo лэрлод леэи ‘впзшиод AOYUR YX ‘по 19,S0g ‘OOZPT $ ‘ON з48м 01 3497] ‘spredog] 115$ JO 190) `00Т “3Iy 178 228 The presence of bright-colored specimens has provided some authors (Satunin, 1914; Bobrinskii, 1944; Novikov, 1956) a basis for suggesting the occurrence or even regular occurrence of Chinese leopards (P. p. chinensis) in the southern Ussuri region. These same two major color types of leopard have also been established for northeastern China (Manchuria) (Baikov, 1927, 1927a, and 1929), where they have been designated distinct subspecies (“‘Manchurian’’ orientalis and ‘‘northern China’’ fontanieri). There is no doubt that these represent only color variants in China, but the boundaries of distribution of the subspecies orientalis in the south are not well defined. Whether there is (or was) in fact an independent form chinensis (fontanieri), described from around Peking, or animals from there belong to orientalis, or are identical with one of the southern forms, or represent a transition from the northern forms to the more southern, is not clear at the present time. Evidently, however, the same form of leopard exists within the Soviet Union as occurs in northeastern China and the Korean Peninsula. 2. Caucasian leopard, P. p. ciscaucasica Satunin, 1914 (syn. saxicolor transcaucasica; the name tulliana is commonly used in Soviet literature for this form). General size, on average, somewhat larger than that of Amur leopard. Color of winter coat very light and pale. Main background color pale, grayish-ocherous, sometimes light gray with sandy or various intensities of reddish, but invariably relatively faint. Color more vivid on back. Sometimes background color even grayish-white, being similar to the main color shade of the ounce. Spots relatively few, usually not pure black, and often with a brownish tinge. Light-colored centers of rosettes usually not darker than main background of coat. Summer coat light, being lighter and less dense than that of Amur leopard. Individual color variability significant; two main color types distinguishable—very light and very dark. Background in light-colored type ordinarily very light grayish-ocherous with a faint reddish tinge; on the back, especially in the anterior part, slightly darker. Sometimes this type is extremely light, i.e., whitish with a light yellow tinge. Most of the spots are compact and relatively small (about 2.0 cm in diameter). Rosettes consist of three to five spots. On the thighs these spots quite often form two bracket-shaped rosettes turned toward each other. Three or four almost complete black rings occur at the end of the tail. Along the middle of the back, in the region of the sacrum, two rows of large (about 4.0 cm long and 2.5 cm wide) elongated spots are present. Background color in dark-colored type ordinarily very dark and reddish. Spots larger (compact ones about 3.0 cm in diameter) and sparser; most of those on the sacrum about 8.0 cm and 4.0 cm in size. A significant number 229 of the rosettes form complete rings. Transverse markings present throughout most of the tail (Flerov, 1935, with modifications).1°° Skull evidently somewhat larger than that of Amur leopards, interorbital region relatively broader (about 22% of condylobasal length), and region of postorbital constriction elongated and lateral sides almost parallel (short post-orbital ‘‘isthmus’’ absent). Nasals relatively moderately narrowed posteriorly and zygomatic arches relatively less curved.!°! Body length 126 to 171 cm (183) and tail length 94 to 116 ст.102 Maximum length of skull in males (six) 202 to 256 mm (M 232.5) and in females (two) 201 to 218 mm; condylobasal length of skull in males (four) 185 to 223 mm (M 206.5) and in females 186 to 188 mm; zygomatic width in males (seven) 133 to 172 mm (M 156.5) and in females 122 to 135 mm; interorbital width in males (four) 41 to 51 mm (M 47.2) and in females 41 mm; width of rostral portion above canines in males (four) 50 to 65 mm (M 58.2) and in females 50 to 53 mm; and length of upper tooth row in males (four) 68 to 75 mm (M 72.0) and in females 64 to 67 mm. Weight may reach 60 kg (Bil’kevich, 1924).103 This subspecies is found in the Great Caucasus, Trans-Caucasus, southern Turkmenia, and southern Tadzhikistan. Outside the Soviet Union Caucasian leopards are found in Iran, Afghanistan, and western Pakistan (Baluchistan). Differences between leopards inhabiting Turkestan and the Caucasus 100Tt has been stated that there are almost no transitional forms between these color types (Flerov, 1935). Such a view is based on a small amount of data. In fact, these represent only extreme forms of a continuous series of variations. It is possible that one or the other type may predominate in different parts of the range. Thus, two color *‘phases”’ of forms pertaining to P. p. saxicolor are shown for northern Iran. In one, distributed in Astrabad and Mezanderan, the main shade is more gray than yellow and spots more dark brown than black; spottiness on trunk shows distinct rosettes; lower portion of flanks, shoulders, and thighs with annular spots; and pattern on head small, mostly rounded spots. More to the east, in Kopet-Dag, Aladag, and the *‘Mastschid’’ [Meshed? — V.H.] mountains, *‘the color of leopards is more yellow than gray. Spots pure black. Annular spots do not form rosettes [1.е., not broken— V.H.]. Lower part of flanks, shoulders, and rear of thighs covered with solid spots. Spottiness on head intense and spots irregular, sometimes angular’’ (T’sukovskii, 1959). Some details of these characteristics as also references to the localization of *‘phases”’ give rise to doubts but on the whole the data are correct. 101 Тре craniologica! features of the subspecies (according to Ognev, 1935; Stroganov, 1962) are only tentative and need revision based on new material. 102 According to data available in literature (Dinnik, 1914; Satunin, 1915; Ognev, 1935). Measurements mainly of skins and stuffed animals (see **Description’’). The biggest leopard measured ‘‘in the flesh’’ had a body length of 171 cm along the curvature of the back, and a tail length of 102 cm (Kopet-Dag:; Bil’kevich, 1924). Craniological measurements are from the same sources and from material of the Zoological Museum, Academy of Sciences and the Zoological Museum, Moscow University. 103See also ‘*Description’’. 180 230 and those of the Far East are factual even though described inadequately (Fig. 101). However, the relations between populations from different parts of the above regions and those of surrounding regions are far from clear. The subspecies name tulliana has long been assigned to leopards of the Trans-Caucasus and Turkestan; this name underscores their identity with the form described in the middle of the last century from the western fringe of Asia Minor. Later (Satunin, 1915), the leopard of the Great Caucasus (P. p. ciscaucasica) was separated from the Trans-Caucasian form based on material from the Kuban Caucasus. Animals living east of the Caspian Sea were usually placed among the Trans-Caucasian forms. This scheme (Satunin, 1914; Ognev, 1935) prevailed until recently, even though the form saxicolor had been described from Astrabad by the end of the 1920’s (Pocock, 1927), which should logically include leopards of Turkmenia, Tadzhikistan, and the Trans-Caucasus. This has been partly suggested by the author’s description who thought that even the Kuban Caucasus leopards belonged to the form захсо[ог.194 The scheme adopted here is that leopards of Turkestan, the Trans- Caucasus, and the Great Caucasus are identical not only with each other, but also with leopards of Iran (at least northern Iran).!°° This single form should bear the less convenient name used here, which has priority over saxicolor. The latter may have importance only when the independence of the subspecies of Great Caucasian leopard has been demonstrated. Such a possibility, however, is extremely unlikely, partly because the animal in the Great Caucasus has been destroyed, or almost so, and is on the verge of extinction. It is difficult to establish the relationship of the leopard population of the above territories with the Asia Minor form tulliana. Evidently, however, it is more distinct than the form designated saxicolor (i.e., ciscaucasica) and the boundary of separation between these forms runs through Mesopotamia (Pocock, 1941).!° In spite of all this, it is quite possible, that leopards of Asia Minor, the Caucasus, Iran, Afghanistan, and Middle Asia represent identical subspecies and hence should be termed tulliana. ш the face of this possibility the recognition of four subspecies 101 Тре form transcaucasica, described by Tsukovskii (1964), is not discussed here. It has not only been separated by upsetting prevailing procedures, but its range, according to the author, is not clear. Thus he exposes his ignorance not only of the distribution and population of leopards in the Caucasus, but also the geography of this country. It is indisputably a synonym for the Middle East form no matter what name is assigned to it. !Fur traders make a clear distinction between Far East and Middle East leopards, but do not differentiate Caucasian from Turkmenian leopards (Kuznetsov, 1941). 16 Тре form tulliana, formerly widely distributed in Asia Minor, is also few in number and its range has shrunk sharply. Its occurrence in the 1940`5 and 1950’s was confined to the extreme southwestern comer of the peninsula between the lower course of the Chediz and Antalya Bay (Kosswig, 1955) or only over a slightly wider area (Kummerloeve, 1967). 179 Fig. 101. Variation in color in Turkmen leopard, Panthera pardus ciscaucasica Satunin. 1— No. 53348, Kharasangli gorge, Kopet-Dag, April, 1948, winter coat; 2—No. 29657, Turkmenia, winter coat; 3 — No. 438920, Sherlauk, Kopet-Dag, October, 1941, winter coat; 4 — No. 29656, Trans-Caucasus, summer coat. Scale same. All skins from collection of the Zoological Museum, Moscow University. Photograph by V.G. Heptner. 181 232 of leopards (saxicolor, sindica, dathei, millardi; Kullmann, 1967—1968) for Afghanistan alone is highly improbable. Apart from light-colored animals of the type ‘‘saxicolor,’’ the occurrence there of the Kashmir form, millardi, is possible, but no other. eta aie $: Outside the Soviet Union many subspecies have been described, numbering over 50 names, and new ones are continually being added. The number of actual subspecies is evidently far less than those recognized by most zoologists (about 25 to 30). Recently, apart from the two forms described above, which are spread far beyond the Soviet Union, the following forms have been recognized, although significant differences of opinion exist in this regard (Haltenorth and Trenze, 1956, with some modifications). Asia: 1) Р. р. minor Hemp. and Ehrenb., 1833——Palestine, Syria, and Arabian Peninsula; 2) Р. р. jarvasi Pocock, 1932—Sinai; 3) P. р. tulliana Valenc., 1857—Asia Minor; 4) Р. р. sindica Pocock, 1930—eastern Baluchistan and Sind; 5) Р. р. millardi Pocock, 1930—Kashmir; 6) Р. р. pernigra Gray, 1856—Nepal and Sikkim; 7) P. p. fusca Meyer, 1794—whole of the Indian Peninsula except Sind, Kashmir, Nepal, and Sikkim; Sri Lanka, Burma, and southwestern China (from Szechwan to Fukien); 8) P. р. delacouri Pocock, 1930—Indochina, Siam, and Malacca; 9) Р. р. melas G. Сиу., 1809—Java and Kangean Islands; and 10) Р. р. Japonensis Gray, 1862—northern (not northeastern! —У.Н.) China. Africa: 11) P. р. panthera Schr., 1777—Morocco, Algiers, Tunisia, Senegambia, and western Sudan; 12) P. p. pardus Linn., 1758—Egypt, eastern Sudan, and to Tanganyika; 13) P. p. antinorii De Beaux, 1923—Fritrea and Ethiopia; 14) Р. р. adusta Pocock, 1927—high mountains of southern Ethiopia; 15) Р. р. brockmani Pocock, 1932—northern (formerly British) Somali; 16) Р. р. nannopardus Thomas, 1904—southern (formerly Italian) Somali; 17) Р. р. adersi Pocock, 1932—Zanzibar; 18) Р. р. chut Heller, 1913—Uganda; 19) Р. р. leopardus Schreb., 1777—forest regions of western Africa from former Spanish Guinea to Nigeria; 20) P. p. reichenovi Cabrera, 1918—steppes of northern Cameroon and the region south of Lake Chad to Congo forests; 21) P. p. iturensis Allen, 1924—forests of Cameroon and Congo; 22) Р. р. shortridgei Pocock, 1932—Angola and Damaraland to Mozambique; 23) Р. р. puella Pocock, 1932—Caocoveld, southwestern Africa; and 24) Р. р. melanotica Ginther, 1885—Cape Colony to southern Rhodesia. The above forms, especially those of Africa, can be combined into groups based on color characteristics and associated with specific natural and geographic conditions. For example, deep-colored animals occur in wet tropical forests and light-colored ones in savannas, while even lighter-colored 233 ones live in desert regions. Quite possibly these groups in fact represent actual subspecies. While some contemporary researchers have a tendency to split groups, others tend to just the opposite. Thus all the four forms of southern Africa (Nos. 19, 22, 23, and 24 of the above list) have been combined into one group and recognized as identical with the Egyptian (nominal) form, i.e., it is assumed that most of Africa is inhabited by a single form (Ellerman, Morrison-Scott and Hayman, 1953). (V.H.) Biology Population. Until the end of the last century leopards were found almost everywhere in the mountains and foothills of the Caucasus, but absent by and large in plains and densely inhabited regions. Leopards were relatively common in the northwestern extremity of the Great Caucasus range. In 1893 they were sighted even at Tuapse and somewhat earlier (in 1875) twice at Sochi (Radde, 1899). From time to time they have been met with north of Fisht and Oshten, not far from Samur station on the Pshekha, a tributary of the Beloe, around Elizavetpol’sk village, and at other places (Dinnik, 1914). In the 1880's lecpards were killed on Dudugush mountain (lower Kishi [пуег]} and occurred in 1895 around Slesarnaya mountain (Shisha River) (Rossikov, 1890; Nasimovich, 1941). In 1896 they were caught on the upper Kishi and they were seen a year later on the upper Sakhrai and on the Pshekisha (Dinnik, 1914). North of Abago (on the upper Beloe) in 1895 N.Ya. Dinnik saw two leopards in the course of one day, and in 1907 in the course of two days thrice heard their roar at midday in the mountains between the Kishi and the Urushten. According to him formerly leopards were caught quite frequently above Psebai in Malyi Laba basin between Umpyr and Zatish’ya settlements. Thus in this region 11 leopards were killed from 1894 to 1896, 2 in 1904, and none in 1905 and 1906. At the end of the last and the beginning of the present century, in the Malyi Laba and Urushten basins, leopards were encountered at many places but evidently confined mainly to the Great and Little Balkhan mountains (Nasimovich, 1941), and in the winter of 1896 to the upper reaches of the Andryuk (Dinnik, 1914). Around 1903 this predator was shot in the Nikits sanctuary in Lysaya mountains. In the Bol’shoi Laba basin all of the old references to the presence of leopards pertain to the region of the Beskes River and Markopidzh mountains. Leopards were sighted quite frequently on Beskes River (Nasimovich, 1941). Around 1905 a leopard was reported as present between the Oshtensk and Guzeripl’ mountains, and some years later near Nagai-Koshka mountain. In 1913 and 1914 a leopard occupied the upper reaches of the Malchepa (Tybga mountain). At the end of the nineteenth and early in the twentieth century in the 234 region described above leopards were subjected to intense destruction by every possible means, including poisoning with strychnine. The destruction of ungulates which serve as food for leopards also played a significant role in the population reduction of this predator. By the beginning of World War I the leopard population in this region had been greatly reduced and in other regions totally eliminated. For example, it had disappeared on the lower reaches of the Kishi and in most sections along the Malyi and Bol’shoi Laba (Nasimovich, 1941). In the years of the Civil War and afterward leopards were sighted once again in some regions where they had previously disappeared. Thus, in 1920 they were seen on the Alousa and near Urushten camp in September, 1932 (Olenich-Gnenenko, 1947; Burchak-Abramovich and Dzhafarov, 1949). Their presence was detected in the Kishi and Beloe basins from 1923 to 1926 in the Chugush, Atamazha, and Dzhemaruk mountains; in the winter of 1933 to 1934 in the Malyi Laba basin; and in 1929 in the Bzyk range. From 1934 to 1936 this predator was sighted several times on the Chugush and Assari; tracks were discovered twice in February, 1934, along the middle Berezova in Chugush region, and in February, 1936, along the Kishi River at Senii camp (1,400 m above sea level) (Nasimovich, 1941). Leopards reportedly lived in the environs of the Black Sea in the past. We can only add here that at the end of the nineteenth century they were not rare in the neighborhood of Veselii village (near Adler); they were also met with in the Bzyk valley and in the Pontic mountains (Dinnik, 1914). During the last 40 years on the Black Sea side of the Great Caucasus range, leopards have been noted in 1926 between the Khosta and Agura Rivers, 4 km from the Black Sea coast; in 1928 at Azhek village (Sochi River), in 1929 two animals caught around Orekhova Polyana (Sochi River). Leopards were seen often in the upper reaches of the Khosta from 1927 to 1929 and also in the upper reaches of the Sochi in Iegosh and Ashako ranges; in the summer of 1934 they were sighted in the Khostinsk branch of the Caucasus preserve. Finally, they occurred in the Tuapse region (Il’minovka village and Yakornaya Shchel’ River). In 1937 leopards were noticed in the territory of the western section of the Caucasus preserve. In 1938 they were encountered a few times in the high mountains on the southern slope of the Great Caucasus range. In 1939 a few appeared within the precincts of Caucasus preserve (Ryabov, 1959). In that same year turs which had fallen prey to leopards were discovered in the upper reaches of the Malyi Laba; the predators still persisted in the early 1940’s to the Dzhumuruk and Dzhugush mountains (Burchak-Abramovich and Dzhafarov, 1949; Ryabov, 1950). From the end of the 1940’s and through 1957 inclusive, leopards were not met with in the Caucasus preserve and apparently had disappeared there (Ryabov, 1959). 184 235 Leopards were observed on the upper Beloe in 1944, on Chugush mountain in August, 1946, and one persisted for quite some time in 1950 around Babuk-aul village in the Lazarev region. In the autumn of 1952 an animal was noted along the Biryuchka River in the Adlerov region. Instances of leopard intrusion into Lazarevskii village near Sochi (Lovetskii, 1956), and in the autumn, 1956, on the upper Bzyk (Ryabov, 1959) have been described. Therefore, in the western Caucasus in the 1930’s leopards were still seen occasionally primarily in two regions—the upper Kishi and Beloe— in the Atamazhi, Tybchi, Dzhamurk, Chugush, and Assara ranges, and the upper Golovinki, Bzyk, Sochi, and Khosty. A few animals lived, apparently, in each region. Later, they became extremely rare here.!° In the last century in the upper part of the Kuban, leopards were frequently killed in the upper reaches of the Urup and Teberda. They have not been noted in recent decades in the Teberda preserve but some were caught in 1958 more south-easterly, in the Kabardino Autonomous Soviet Socialist Republic, on the upper Chegem, a tributary of the Terek (Alekseev, 1958). In the former Terek and Dagestan districts leopards have been reported Fig. 102. Habitat of leopards in central Kopet-Dag. Biotope of goat (Capra aegagrus — left slope) and mountain sheep (Ovis ammon cycloceros—more gentle slopes on right). May, 1963. Photograph by М.1. Ishadov. ‘07H owever, it was later reported that stray encounters at Babuk-aul (forest preserve) have taken place (Aleksandrov, 1965). (V.H.) 183 236 at many places but comparatively rarely, being common only in the upper Avarsk Koisa. One leopard was caught not far from Evdokimov fortress on the Argun River (Dinnik, 1914). In the spring of 1924 another was killed at Agach-aul near Makhachkala city (Burchak-Abramovich and Dzhafarov, 1949). In Dagestan at the beginning of the present century the leopard was encountered more or less regularly only in the region of the upper Andiisk and Avarsk Koisa; the animal was apparently already absent in the central Caucasus (Heptner and Formozov, 1941). On the southern slopes of the central parts of the Great Caucasus range leopard was earlier generally common in the Alazan valley, and was killed many times around Zakatal, Lagodekh, Nukhu, and the Bumsk gorge. In December, 1931 one was killed in the Zakatal’sk preserve, and in November, 1937 at the pass opposite the city of Zakatal (Markov and Mlokosevich, 1935; Vereshchagin, 1942). In recent years none have been seen in the ” Zakatal’sk and Lagodekh preserves, but tracks were encountered in 1937 to the east, on the upper Tala-chai. All the information given above suggests that leopards are now met with extremely rarely in the Caucasus proper. In the western Trans-Caucasus and also in the coastal region reports of leopards were very rare even in the last century. Some animals were noted between Kur’ and Rioni. In the Borzhomi region three were bagged over several years. Near Tiflis [Tblisi] one was killed in the early 1850's and another caught in the 1950’s (A.A. Nasimovich). Leopards were also not rare in the Karabakh and Talysh ranges (Radde, 1899; Dinnik, 1914). In the Talysh, for example, G. Radde was given 12 fresh skins in 7 weeks in 1866. Over the last 40 years in the Trans-Caucasus leopards have become extremely rare, and occur irregularly. They were still encountered fairly regularly in Talysh and Zangezur ranges, although already rare. In this period they were noted at the following places: 1929 (or 1930) near Kharmandal village on the Mugana (dead female; Sokolov, 1931); 1939 between Shangolu and Karabakhlyar (Sarai-Bulagsk range); 1940’s in Krasnovo Kirovobad region; 1946 in Kul’badzhar region; 1946 in the Zangezur range east of the city of Ordubad (two leopards killed); 1947 in Shikh-Yurda region on the western slope of the Zangezur range; and 1947 on Ilanludag mountain and sometimes Darrydag and Megrinsk mountains (Burchak-Abramovich and Dzhafarov, 1949). In 1930 a leopard was caught at Ogra in the Lenkoran region and another animal killed there on March 13, 1930 (Starchikhin, 1930); in 1944 a leopard was caught 30 km from Lenkoran and in 1946 a juvenile was killed on the Talysh (Alekperov, 1947). In 1952 an animal was caught in the hills near Araks gorge at Neram station (Sadykov, 1952). In 1953 a leopard was killed in the Zangezur range near Bartsruni in the 25 Azizbekov region, and in the Ararat valley near Dvin village in the Artashats region; in 1956 one was caught roughiy 25 km east of Araks in the Vedinsk region in Khosrov forest where it had lived for several months (Geilikman, 1956). In recent years leopards have been encountered from time to time in southern Armenia in the Kafan region. In 1957 two leopards were caught in Armenia, and another in the winter of 1959 (Zaritap village in the Azizbekov region; Mnatsakanyan, 1959). In southern Armenia in the 1950’s on the average one leopard was killed every one to two years (Gambaryan, 1957). In 1964 one animal was confined to the Shamshadinsk region (Kasumov, 1964). In the mid-1960’s in Armenia leopards were still encountered in the Zangezur range but were very rare; the population there was supplemented by animals coming from Iran. In the rest of the Republic this species was exceptionally rare and reported only as an intruder (V.G. Heptner). In Georgia at Akhaltsikhe a hunter killed three leopards in the winter of 1959 to 1960 (Galandyan, 1960). The animals had evidently arrived from Turkey. In Azerbaidzhan in 1958 a leopard was caught in the Gaidag gorge (Khromov, 1958) and a second killed near Lenkoran in the winter of 1958/59. Fig. 103. Steppe-covered plateau in the high mountains of the central Kopet-Dag — pasture of mountain sheep (Ovis ammon cycloceros) and site of residence and hunting of leopards. In the center is an old juniper tree (arborescent juniper shrub), a shady refuge for sheep and leopard. April, 1963. Photograph by N.I. Ishadov. 184 238 In December, 1959 one was killed near Atrek village, Mardakerts region of Nagorno-Karabakh Autonomous District. In this republic one of the last leopards was killed in January, 1967 in the Gadruts region near Tug village (Aslanov). To what extent leopards are rare in the Caucasus and in Trans-Caucasus can be judged from the fact that skins come singly for tanning. For example, only one skin was prepared in Azerbaidzhan in 1937 and 1938. In the preceding decade for the whole of the Caucasus, only 10 leopards were caught, most of which were killed in the Zangezur range and on the Talysh (Vereshchagin, 1947). In 1960 on the Talysh four hunters bagged only five leopards and a minimum of nine were killed throughout Azerbaidzhan from 1960 through 1964 (A.A. Sludskii). Only two to four animals have been killed per annum in this republic since then (Aliev and Nasibov, 1966). In the Trans-Caucasus leopards were encountered from time to time even on the plains, especially on the Kurinsk lowlands; one was killed on the steppe close to Salyany (Radde, 1899). On February 20, 1946, on the northern Apsheron Peninsula at Bil’gya village, 25 km north of Baku, an animal arriving at the time of a snowstorm fell into a well (Dzhafarov, 1946; Alekperov, 1947). In the Caucasus and Trans-Caucasus leopards disappeared rapidly as a result of ruthless killing, cultivation, and a reduction in population of ungulates. In Armenia mountain goats and sheep, which live in association with leopards, were almost extinct even in the 1960’s (V.G. Heptner). Leopards will evidently disappear in the Caucasus in the near future. Early in the present century leopards were ‘‘дайе common’’ at Kopet- Dag in Turkmenia. ‘шт 1911 I heard about panthers everywhere and almost daily saw their tracks and lairs in the sands of high montane streams and also spoor’’ (Dinnik, 1914). In the first two decades of the twentieth century in Ashkhabad there were hunters who had each killed several leopards (V.G. Heptner). In the 1920’s it was thought that the average density was 0.25 animals per 100 km? throughout the whole of the range in this republic. Even in the 1940’s and 1950’s, when mountain sheep were abundant everywhere in the Kopet-Dag, especially in the higher regions, leopards were common throughout the range in spite of a lower population than in the 1920’s. In the Kopet-Dag some 15 leopards were killed annually in 1940 and 1941 (V.G. Heptner). Leopards were very common in the mountains between Kushka and Murgab (Chengurek mountains). They were particularly numerous west of Badkhyz in the Gyaz’-Gyadyk mountains, an area extremely rich in argali sheep in those years. Leopards were regularly found and killed there, at the rate of 14 animals per year in 1947 and 1948, in a very small area (about 500 km). The leopard population in the Gyaz’- Gyadyk was larger than that found anywhere else in the Soviet Union. Apart Fig. 104. Broad valley with a spring, middle belt of the central Kopet-Dag, with juniper, mountain maple, honeysuckle, small fruit cherry, and other vegetation, used as a water hole by mountain sheep (Ovis ammon cycloceros) and goats (Capra aegagrus), and a source of water and prey for leopard. August, 1963. Photograph by N.I. Ishadov. from the abundance of argali sheep, this was due to the arrival of migrant animals from Iran. In the 1960’s the population shrank drastically, in part due to a reduction in number of mountain sheep (V.G. Heptner). Leopards were common in the Great Balkhan but due to ruthless killing and a sharp reduction in sheep they disappeared or became very rare in the 1950’s. They were always rare in the Little Balkhan and long absent in the Kyuren’-Dag, where they probably had disappeared even in the last century (V.G. Heptner). In Turkmenia 19 (1959*) to 41 (1956) skins were tanned annually in the 1950’s.!9 Leopards were very rare in southwestern Tadzhikistan and southern Uzbekistan (Babatag, Aktau, Karatau ranges, and others). In southern and southeastern Trans-Baikal in the last as well as the present century, leopards were sighted rarely and very rarely intruded into the central Amur region. *Listed in ascending order of skin tanned— Suppl. Scient. Edit. 108These skins might also include those of cheetah. Some leopard skins were no doubt retained by hunters. 186 240 In the last century in the Far East, in the Ussuri territory leopards were rare. They were found often only in regions adjoining China. In 1867 during a six-month journey throughout the Ussuri territory, covering a total of 2,250 km (partly by boat), N.M. Przheval’skii (1870) discovered only two sets of leopard tracks and obtained a single skin with much difficulty. Much of his route lay along the coast of the Sea of Japan, covering places where leopards were reportedly often encountered. Leopards were rare in the Far East even in the early years of the present century. In 1912 two animals were killed in the Amur region, and eleven in the Primorsk district (Baikov, 1927a). According to other data, however, only - one or two were caught annually in these regions between 1911 and 1914 (Tselishchev, 1924). In the 1920’s two or three animals were caught per year in the Ussuri territory (Solov’ev, 1925). Leopards were even rarer in the 1950’s and 1960’s. However, encounters were more frequent due to intensive cultivation of formerly virgin areas and human population growth. As in the last century, this predator is now “‘not a rarity’’ along the western coast of the Amur Gulf. Some 15 to 20 leopards presently live in the montane forest region lying to the north of Suifun River and thence southwest to the state frontiers with the Korean People’s Democratic Republic and the Chinese People’s Republic (Vasil’ev et al., 1965). According to other data this number represents the leopard population inhabiting the whole of the Primor’e (С.Е. Bromlei).!° In “‘Kedrovaya Pad’ ’’ preserve (southwest of Vladivostok) four to six leopards were regularly seen in an area of 9,500 hectares in 1940 and 1941 (Mirolyubov, 1941); a family of two to four animals lived there т a section 40 km? in the mid-1960’s (Vasil’ev et al., 1965). The high population density of leopards in this preserve is explained by the abundance of ungulates there. _ Habitat. In the Caucasus leopards inhabit subalpine steppe-covered meadows, deciduous forests, and dense shrubs; they are generally confined to rock outcrops and talus slopes. The main conditions determining their habitat are a high population of bezoar goats, turs, roe deer, chamois, deer, and wild pigs, and the availability of sections with shallow snow in winter. Thus, in the western Caucasus leopards live high in the hills and are seen quite often in summer in the alpine zone and even at permanent snow line at a height of 2,600 to 2,700 m above sea level, and sometimes even higher; in the central part of the range they ascend to 3,000 to 3,500 m above sea level (Dinnik, 1914). They are confined in the latter area to a fairly high level even in winter and have been sighted at 1,500 m (February, 1934; Berezova River) and 1,400 m above sea level (February, 1936; Kishi River). 1091 1970, 32 to 35 leopards were counted in Ussuri territory (S.P. Kucherenko). (V.H.) 187 188 Fig. 105. Steppe-covered plateau with juniper trees (arborescent juniper shrubs) and limestone cliffs at the top of the Great Balkhan mountains, a place formerly very rich in mountain sheep (Ovis ammon cycloceros— plateau) and goats (Capra aegagrus — cliffs). Leopards were also common there. About 2,000 m above sea level. February, 1963. Photograph by E.N. Matyushkin. On both occasions they were close to the wintering sites of chamois. The depth of the snow cover ranged from 70 to 100 cm (A.A. Nasimovich). These mountains are rich in turs, chamois, and at times deer. On the Black Sea side of the western Caucasus leopards often do not reach the alpine zone, characterized by extreme rockiness, slopes overgrown with dense cherry laurel, azalea, Pontic rhododendron, and Caucasian bilberry. Many areas of these mountains are almost impenetrable to human. Chamois are quite common in the higher levels and roe deer and wild pigs also occur at sui- table places. In these mountains leopards range from the sea coast up to the mountain tops. In the Trans-Caucasus this predator lives (lived) in the treeless or nearly treeless rocky mountains and gorges where bezoar goats and Trans-Caucasian mountain sheep are abundant. In southern Armenia it is confined to 550 to 950 m above sea level on slopes with steppe vegetation and rarely ventures into forest regions (Dal’, 1954). Leopards sometimes live even in the plains, for example in the Lenkoran lowlands, if they are covered with dense forests or reeds, prickly shrubs, and various kinds of lianas, and inhabited by wild ine) aS iw) 187 Fig. 106. Gorge dissecting a plateau summit. Left, an old juniper tree (arborescent juniper shrub). Typical leopard habitat in the Great Balkhan mountains. February, 1963. Photograph by E.N. Matyushkin. pigs. On the level steppes leopards occur only as intruders. In the Kurinsk lowlands they are encountered from time to time in reed-cattail thickets around water reservoirs, and also in tugais along the Kur’ and Araks (N.K. Vereshchagin). In Turkmenia leopards mainly inhabit mountains featuring rocks and gorges, and almost devoid of trees, or sparsely covered with juniper and pistachio. Along the cliff slopes of the gorges of these mountains and uplands, tangles of dog rose, blackberry, barberry, and in places native pistachio, are encountered, which grow on the terraces, “‘shelves’’, and in their bottoms. Leopards are confined here to sites rich in bezoar goats and Turkmenian mountain sheep (Ovis ammon cycloceros) or only the latter; the predator ascends in summer to the uppermost levels and descends in winter to lower levels following goats and sheep. The Gyaz’-Gyadyk mountains in southern Turkmenia (Badkhyz), where leopards are still relatively common, are not high, but are interrupted by deep gorges with steep slopes, and limestone and sandstone outcrops. bo > w 188 189 Fig. 107. Slopes and small valley with pistachio thickets and rich herbaceous cover of sagebrush, meadow grass, and giant umbellifers in Gyaz’-Gyadyk. Characteristic biotope of argali sheep (Ovis ammon cycloceros), and leopards found in association with them. Pistachio represents a favorite shady refuge of both species. Badkhyz preserve. October, 1959. Photograph by A.A. Sludskii. Mountain slopes are covered with sedges, meadow grass, and forbs while along the range, its slopes to the very bottom are covered with perennial trees of the true pistachio, the diameter of which at ground level may reach 1.0 m. Sometimes these trees form characteristic bushes. They are well separated and do not form close clusters (“‘Pistachio savanna’’). Even old trees are not tall but very dense; their crowns form a semicircular dark green marquee since their low branches almost reach the ground. - At the end of summer, when pistachio fruits ripen, ungulates congregate there; wild pigs, Turkmenian sheep, goitered gazelles, and porcupines. Leopards are attracted by the abundance of prey. Goitered gazelles are particularly numerous around pistachio trees in arid years. Along the bottom of gorges, usually overgrown with wild figs, small saline springs occur, for example at Kerlek, containing 8.0 to 20.0 g salts per liter. Reeds grow in a narrow belt along some of these springs and wild pigs inhabit them. Kulan, wild sheep, sometimes goitered gazelle, and various carnivores—wolf, hyaena, and cats — gather at the more accessible springs in summer. Leopards also drink from such springs. Predators lie in wait for ungulates at these watering places. In the Badkhyz leopards have been found in the deep Er- 189 Fig. 108. Spring in one of the gorges of the Gyaz’-Gyadyk mountains serving as a watering site and hunting ground for leopards. Slope streaked with trails of ungulates coming to water, mainly mountain sheep, also wild pigs, sometimes goitered gazelles and kulans. Badkhyz preserve, southern Turkmenia. February, 1965. Photograph by V.G. Heptner. oilan-duz depression, hunting for wild sheep among chinks [arroyos | over- grown with herbs and shrubs such as saltwort, locoweed, and others. Along these cliffs sheep from Gyaz’-Gyadyk, trailed by leopards, enter the very heart of the desert at Dyzyl-Dzhar where, in addition to sheep, goitered gazelle, some wild pigs, and kulan are common visitors (Heptner, 1956; A.A. Sludskii). In the Atrek basin in Kopet-Dag (Sumbar and Chandyr Rivers), 190 192 Fig. 109. Spring at site of Kerlek, watering place of leopards in Gyaz’-Gyadyk mountains. Badkhyz preserve, southern Turkmenia. May, 1962. Photograph by A.A. Sludskii. leopards also live in large gorges with giant nut-bearing trees, thickets of fig, and other trees entwined with vines, impenetrable tangles of blackberry, etc. (Ai-Dere, Pop-Dere, etc.) (V.G. Heptner). The leopard’s range in Turkmenia almost totally coincides with that of Turkmenian sheep. This carnivore is quite common in regions where bezoar goats are absent or rarely encountered; it is also absent at places where goitered gazelle are numerous, but is present even in unfavorable deserts if they are inhabited by sheep (Badkhyz). On moving into a desert the carnivore confines itself to cliffs where it can find shelter (Heptner, 1956). In Tadzhikistan and Turkmenia leopards live in low mountains covered with desert vegetation and normally do not ascend higher than 1,500 m above sea level. In Babatag (Uzbekistan) leopards keep to deep and narrow gorges and precipitous rocky slopes which alternate with relatively gentle slopes covered with pistachio shrubs. At the bottom of these gorges many fresh or saline springs are present, and tamarisk, poplar, and sometimes reeds, ambary, and grapevines are relatively abundant. Ironwood, almond bushes, and Persian ironwood grow on the steep rocky slopes near such places. Mountain sheep and markhor, the main prey of leopards, live on the slopes of these gorges and in the watersheds between them. Leopards in Babatag 190 246 — Fig. 110. Saline lakes, buttes, and chink wall (right— habitat of mountain sheep and leopards. Hyaena also live here. Er-oilan-duz depression in Badkhyz desert, southern Turkmenia. May, 1962. Photograph by A.A. Sludskii. live at heights of 600 to 900 m above sea level (Bogdanov, 1952; Ishunin, 1961). In the Far East, in Primor’e, leopards are confined to a narrow coastal - strip with Manchurian type broad-leaved forests and at places with a rather low snow cover. They visit rocky sections along the sea coast very rarely, on the average two or three days in a year. For example, on Tuman mountain leopards were sighted only in March, 1942; December, 1948 [sic]; and January, 1946. Between 1944 and 1946 some stray tracks of leopards were detected on two occasions in autumn along the coast extending from Valentine Bay to Preobrazheniya Bay (Bromlei, 1963). The animal has been sighted on occasion, mainly in winter, deep inside the mountains, on their western slopes. Evidently the high snow cover of these areas poses a serious obstacle to movement. On Cape Gamov, when the snow cover is high, leopards find movement very difficult and prefer confinement to one place on talus slopes (V.V., 1927). In winter, during wanderings, they keep to snowlree rocky slopes facing south. Rocky sections are favored over adjacent sections devoid of rocks. This carnivore generally avoids roads, human and animal trails, frozen ice crusts, and rivers; however, it takes advantage of deer trails for hunting in certain areas. 247 191 Fig. 111. Buttes of the Er-oilan-duz depression; northern chink visible in background and bottom covered with solonetzes and saxaul shrubs. Habitat of argali sheep, goitered gazelle, kulan, leopard, and hyaena. June, 1948. Photograph by V.G. Heptner. 191 Fig. 112. Gigantic argillaceous cliffs (‘sheep cliffs’’) in Kyzyl-Dzhar gorge dissecting Badkhyz desert plateau. Habitat of arkhar sheep, visited by leopards. Southern Turkmenia. February, 1961. Photograph by M.V. Heptner. 248 19 > a Fig. 113. Forest of *‘Kedrovaya Pad’ ’’ preserve. Site of permanent residence of leopards. Tiger, leopard cat, sika deer, etc. also live here. June, 1958. Photograph by A.G. Pankrat’ev. In the Far East leopards live not only in montane forest regions but also in valleys and plains. For example, at the end of the nineteenth century, these animals were met with in meadowlands in the Pre-Khanka lowland. Nowadays none are found in the lowland meadow-forest sections of the Mo and Ге interfluve ([Гаке] Khanka tributaries) of the Pre-Khanka lowland. South of Lake Khanka in 1956 to 1961 these predators lived in rocky creeks and among reeds in the floodplains of the Chapigos valley. In the 1950’s they were seen from time to time in meadow-forest sections in the floodplains of the Amur and Bir (Rakov, 1965). Their distribution in the Far East is usually associated with the presence of sika and roe deer. It is quite possible that in Primor’e this species is only an intruder and hence breeding there would be merely incidental (G.F. Bromlei). Food. Wild ungulates —bezoar goats, Низ, mountain sheep, chamois, roe deer, deer, and wild pigs—constitute the main prey of leopards in the Caucasus and Trans-Caucasus. Sometimes they also catch European hare, pheasant, rock partridge, Caucasian black grouse, snowcock, and porcupines (in Talysh) (Dinnik, 1914; Vereshchagin, 1942). At places where ungulates are abundant, for example in the former Kuban district, leopards almost do 193 194 Fig. 114. Deer Cliff in “‘Kedrovaya Pad’ °° preserve near Vladivostok. Regular favorite habitat of leopards. June, 1964. Photograph by A.G. Pankrat’ev. not attack domestic animals, but in the Trans-Caucasus attacks on various domestic animals, including dogs and poultry, have been reported frequently. Bones of horses, asses, cattle, wild goats, and birds have been found around a leopard’s lair in the Zangezur range near Kalita village (Burchak- Abramovich and Dzhafarov, 1949). In Turkmenia, in the Kopet-Dag, leopards hunt mainly bezoar goats and Turkmenian mountain sheep, and more rarely wild pigs and domestic animals such as cattle, horses, asses, sheep, goats, and dogs. Not only the young but even the adults of these animals are attacked (Dinnik, 1914; Bil’kevich, 1924; Morits, 1935). Around the lair of a leopard in the Gyaz’- Gyadyk (Akar Cheshma) there were many bones of goitered gazelle (Filippov, 1956). The main prey of leopards are Kopet-Dag sheep in most regions of Turkmenia, especially in Badkhyz (Heptner, 1956). Sometimes leopards attack porcupines but pay the price. In the soles of one leopard’s paw were found nine pieces of porcupine quills measuring up to 1.0 cm. Necrosis had set in around the pierced areas (5.1. Bil’kevich). In February, 1951, in Badkhyz (Gyaz’-Gyadyk, Kepel village) a leopard was killed which had many pieces of porcupine quills in its paws (Rustamov and Shcherbina, 195 250 1957). Another one caught on August 4, 1960, in Badkhyz had quills up to 4.0 cm in its limbs and other body parts (Gorelov, 1963). In March, 1955, a leopard was located in the burrow of a porcupine which it had killed (Dement’ev and Rustamov, 1956). Leopards take to attacking domestic cattle mainly in snowy winters or when the population of wild ungulates has shrunk. For example, in the extremely snowy winter of 1921 an animal lived quite close to a village and daily stole a sheep (Bil’kevich, 1924). The reduction in number of bezoar goats and Turkmenian sheep observed in recent decades in Kopet-Dag has Fig. 115. Leopard habitat in ‘‘Kedrovaya Pad’ ’’ preserve, near Vladivostok. August, 1969. Photograph by G.N. Simkin. 195 196 251 led to an increase in attacks by leopards on domestic animals in that region. Some feed on cattle fairly regularly, and are termed “‘village panthers’’ in India. As late as the spring of 1951 in the Kyzyl-Arvats region three leopards were killed when they attacked cattle. One animal regularly attacked horses and killed 9 in 1949, 17 in 1950, and 3 in the first three months of 1951. To safeguard them against leopard attacks horses in a collective farm had to be driven to another region. A leopard killed a horse even in the Badkhyz preserve (Dement’ev and Rustamov, 1956). In Dmitrievka village, 50 km from Goek- Tepe, the predator caught nearly every dog available (Dinnik, 1914). In the Ussuri territory the main prey of leopards are roe deer, sika deer, Manchurian wapiti, musk deer, moose, and wild pigs; more rarely the predator catches hare, badger, fowl, and even ‘‘mice’’ (Baikov, 1915 and 1927; Yankovskii, 1882). In ‘‘Kedrovaya Pad’ ’’ preserve (southwest of Vladivostok) roe deer is the main prey of leopards year-round (Vasil’ev et al., 1965). In the southern Ussuri territory (Tumannaya knoll) it catches hares and from time to time gorals (С.Е. Bromlei). In ‘““Kedrovaya Pad’ ”’ preserve leopards attack raccoon dogs, often abandoning them; they prey on young, Eurasian black bears (Ursus tibetanus) less than two years old (Vasil’ev et al., 1965). These bears are preyed upon even in the north, in the Pri-Amur (Sysoev, 1966). In Primor’e, leopards attack sika deer held in parks (Menard, 1930; С.Е. Bromlei). In the forest nursery at Pushkino a leopard killed in one day three deer, two the next day, and so on. Because of the extreme rarity of leopards in Sudzukhin preserve, between 1936 and 1948 not a single mortality of sika deer was attributable to this predator (Bromlei, 1956). The stomach of a large male caught on August 4, 1960 at Badkhyz contained 2.0 kg of meat from a mountain sheep (Gorelov, 1963). In zoological gardens leopards are fed daily about 3.0 kg of meat (Obudhova and Shakhnazarov, 1949). The report that a hungry predator can consume almost an entire roe deer in a single day (Baikov, 1927) is clearly an exaggeration. In China leopards catch deer, wild pigs, gorals, monkeys, hares, large birds, and domestic animals, particularly dogs and pigs (Shou, 1958). In India this predator hunts for sambar deer, wild pigs, jackals, monkeys, hares, and porcupines, and also eats lizards, snakes, and crabs. In Kashmir it attacks hangul deer when their movement is impeded by deep snow. Leopards often prowl near villages and sometimes become a pest of domestic cattle, destroying calves, asses, horses, goats, sheep, and other animals, but most often dogs, for which they will even break into a house. Jackals and dogs represent their favorite prey. They also eat carrion irrespective of the degree of decomposition. In Africa the biggest animals attacked by leopards are camels, cows, kudu antelope, and asses. The predator usually feeds on 195 iw) Cn Ww Fig. 116. Site of transient and resident leopards in Sudzukhin preserve on the eastern slope of Ta-Chindzhan range. Southern Ussuri territory. August, 1958. Photograph by V.E. Prisyazhnyuk. moderate-sized and small animals such as small antelopes, wart hogs, sheep, goats, pigs, aardvarks, monkeys (baboons), porcupines, domestic fowl, and small rodents; it also attacks jackals, foxes, genets, and domestic dogs. Leopards sometimes attack man and some become man-eaters, i.e., man may serve as their main food item (see ‘‘Practical Significance’’). Home range. Not much is known about this. Zoologists are of the opinion that, outside the breeding season, leopards prowl continually in search of sites 196 197 Fig. 117. Montane mixed broad-leaved forest with an admixture of fir and Korean nut-pines. Habitat of leopards in ‘*Kedrovaya Pad’ ’’ preserve near Vladivostok. December, 1962. Photograph by A.G. Pankrat’ev. abundant in wild animals. This view is obviously only partly justified. At places where wild animals are abundant leopards live permanently or perform only vertical migrations, trailing herds of ungulates and avoiding snow. A family of two to four leopards lives regularly in “‘Kedrovaya Pad’ ”” preserve in Primor’e. Year after year they have occupied the same section and rarely gone outside the preserve boundaries. This section covers the upper reaches of the Kedrovaya River where forests are dense and narrow mountain crests and peaks with rocks and talus slopes stand out prominently. The area of this section is about 40 km? and the leopards have made permanent trails within its confines (Vasil’ev et al., 1965). Two or three leopards lived permanently in the Chapigou River valley of Primor’e from 1956 through 1961 and possibly constituted a single family. These animals were seen in an oval-shaped hunting territory, the longer side of which was about 40 km. Annually, after the commencement of frosts, the predators descended from talus slopes into valley sections of reeds housing abundant roe deer and raccoon dogs. In 1960 and 1961 a lone leopard on the southern slope of the Pidan range occupied a hunting region 30 to 197 254 Fig. 118. Mixed coniferous—broad-leaved forest in **‘Kedrovaya Pad’ ^` preserve near Vladivostok. Habitat of leopards. Spiny actinidia vine on Korean nut-pine visible in foreground. October, 1965. Photograph by A.G. Pankrat’ev. 35 km long and 10 to 15 km wide and fed mainly on roe deer (Rakoy, 1956). A leopard remained close to a deer park for several years and eventually had to be put down (Menard, 1930). In the Caucasus in recent decades the individual territories of leopards were very large, occupying as much as 100 km? (Nasimovich, 1952). Burrows and shelters. The den in which a leopard raises her cubs is set up in a cave under an overhanging rock or under a wind-felled tree, 199 198 255 located usually in a particularly dense, inaccessible section of a gorge. А lair in the Kishi valley (western Caucasus) was built on the ground in a wide patch of uprooted trees. The ground at the site of the den and around it had been churned and the grass well beaten. Feces of the female and cubs and bones of various ungulates were scattered around the den. A second . den in the same patch of wind-felled trees had been set up under a rock (Dinnik, 1914). In Turkmenia a leopard was caught in the burrow of a porcupine (see above). A female with two cubs was found in the burrow of this rodent near Akar-Cheshma. Around the burrow were many bones of goitered gazelle (Filippov, 1936). In northeastern China shelters for raising cubs are located among dense shrubs, near talus slopes on the southern slopes of hills, in caves, and under overhanging rocks. The den [floor] is covered with dry leaves and grass prior to parturition (Baikov, 1915). If a den with very small cubs is threatened, the mother carries them in her teeth, one at a time, to a new site. Males and lone females also rest during the day in caves, rock fissures, or dense thickets. In Turkmenia caves and very deep niches in rocks usually serve as leopard shelters (V.G. Heptner). A leopard shelters on warm summer days in the shade of an overhanging rock but loves to sun itself on an exposed rock in cold weather. Sometimes the predator sleeps during the day in well- Fig. 119. Site of permanent occupation of leopards in ‘‘Kedrovaya Pad’ ”’ preserve, near Vladivostok. September, 1970. Photograph by V.G. Heptner. 256 protected places such as open sites on slopes, behind rocks on ledges, etc. (V.G. Heptner). In India, as well as in Turkestan, the den is sometimes made in a porcupine burrow (Pocock, 1939). Daily activity and behavior. Leopards set out to hunt usually an hour or two before sunset, hunting through the first half of the night, and again in early morning. In spite of the fact that a leopard is essentially a crepuscular animal, from time to time it chases prey even during the day, especially on cloudy, cold days and in winter. It goes to water holes with the onset of twilight, but sometimes even during the day. The leopard is a cautious, secretive animal, but never timid. On encountering a man, it does not attempt to hide quickly as do many animals, but withdraws unhurriedly and with no exhibition of fear. Sometimes it does not run away even when a shot has misfired. Instances are known when leopards, in sight of hunters, carried off a goat wounded by them, or of pursuing game unmindful of the shouts of the people or their shots. Unlike ounce and cheetah, an injured leopard often attacks the hunter or chases 198 Fig. 120. Broad-leaved forest with an admixture of nut pines and fir. Biotope of Яя leopards. ‘‘Kedrovaya Pad preserve, near Vladivostok. September, 1970. Photograph by V.G. Heptner. 257 him. Hence т most southern Asian countries leopard hunting is considered extremely dangerous. Elsewhere, instances have been recorded of an animal wounding several men who set out to catch the predator. When attacking aman a leopard rears up on its hind legs and strikes him with its forepaws or attempts to grab him by the throat. A leopard can conceal itself exceedingly well and often remain within just 10 m of men passing by. It climbs well and in rescuing itself from pursuit generally climbs inaccessibie rocks or trees and conceals itself, making detection very difficult. Its every movement is graceful, supple, and agile. When a leopard moves, it progresses as quietly and lightly as a small cat. This predator usually avoids water and prefers to cross small brooks on fallen trees or a bridge. On occasion it goes into water and swims across large rivers. In the western Caucasus in February one year a leopard crossed a fiord through an unfrozen rivulet, the depth of which was 50 cm (A.A. Nasimovich). Vision and audition are excellent in leopards but olfaction poor, due to which these animals do not hunt by scent. They employ various methods for hunting. Usually the predator will lie in wait for prey, hiding in bushes among rocks, on an outcrop or tree near a trail, water hole, or solonetzes and attack the unsuspecting victim from an ambush in a single leap of 6.0 to 8.0 m. On spotting the prey the predator conceals itself and crawls forward 50 quietly that detection is extremely difficult. In winter, on a snowfree slope with a background of dried leaves or grass, it is almost impossible to locate a concealed leopard even at a distance of 50 т. In grass 50 cm high, when the predator starts to creep, it completely disappears from view. When the quarry is close by, the leopard chases it in long bounds. Sometimes it sprints after the quarry, executing bounds of 7.0 to 8.0 m length so rapidly that it can quickly overtake a roe deer, goral, musk deer, bezoar goat, or dog. Such chases of deer, roe deer, and chamois have been observed in the Caucasus and chases of bezoar goats seen in Kopet-Dag. Leopards are usually capable of overtaking quarry over a distance of not more than 40 to 50 m. In ‘“‘Kedrovaya Pad’ ”’ preserve in Primor’e, in winter the leopard conceals itself to catch roe deer and creeps forward from the leeward side of the prey in the late morning hours when, after grazing, deer climb the slopes toward their beds. Sometimes a leopard catches one of them by hiding under coastal cliffs (Vasil’ev et al., 1965). In-attacking large animals a leopard generally rears up on its hind legs and inflicts severe blows with its forepaws. In the case of small animals it twists the neck in one stroke and then bites the skull. Larger animals are first knocked down, after which the predator clamps its jaws on the neck at the occiput or rips open the throat. A female sheep in Kopet-Dag killed 200 by a leopard had her throat slit, withers broken, and wide gaping wounds 258 on the muzzle inflicted by the claws; the bone under the eyes was evidently pierced by the large claws of the forepaws (Morits, 1935). In killing a porcupine the predator grabs it from the front just behind the head, thus protecting itself from the quills (Corbett, 1957). On entering an area rich in game, a leopard usually kills a single animal and does not frighten the rest. In this respect it is similar to tiger but differs from wolf, which usually attempts to wound several animals and thus drives them away. Even within a deer park a leopard usually kills only one deer at a time. In the Ussuri territory the leopard hides the carcass of a roe deer among the roots of upturned trees or under wind-felled trees, feeds on it for two to four days, and hides nearby (Vasil’ev et al., 1965). The predator sometimes conceals the remains of its prey in hollows and rocks, or in the tropics, in trees. In Kopet-Dag the leopard usually returns to any unfinished prey. Quite often it rests during the day near the carcass of a killed animal and drives off vultures. These birds, numerous in this region, can pick clean the carcass of an argali sheep within an hour. Evidently to safeguard its prey, it will jump to a height of several meters, springing into a juniper tree with the quarry in its mouth. In spite of its comparatively small size, the leopard possesses tremendous strength. Once this predator was seen carrying a young horse it had killed over rocks for a distance of 50 m. On another occasion a leopard jumped onto a rock 3.0 m high with a mountain sheep clamped between its teeth. Ascending upward along a steep slope, this cat can carry a female bezoar goat rather effortlessly (V.G. Heptner). Seasonal migrations and transgressions. As noted above, outside the breeding season when food is inadequate or access to it difficult (deep snow cover, thin snow crust), leopards migrate from one area to another. Fairly regular vertical migrations of this animal have been observed in the Caucasus where, depending on snow thaw in spring, the predator ascends to the alpine zone, trailing ungulates, and descends with them in autumn. Normally an inhabitant of the mountains, it sometimes migrates far into the plains (100 km or more). 5 In the Caucasus, in Dagestan, in the early part of the present century leopards, though very rare, were nonetheless sighted fairly regularly in the upper reaches of the Andiisk and Avarsk Koksa, and in the spring of 1924 caught at Agach-aul near Makhachkala (Burchak-Abramovich and Dzhafarov, 1949). On the Black Sea coast leopards were sighted often at the end of the last century, and in the 1920’s to 1930’s, at Adler, Khosta, and Sochi. They were found there even later. In 1950, for example, a leopard resided for quite some time near Babuk-aul village in the Lazarev region and in the autumn of 1952 one was noted on the Biryuchka River at Lazarev village near Sochi (Lovetskii, 1956). 03 259 In the Trans-Caucasus leopards were sighted fairly regularly in the present century in the Talysh‘and Zangezur ranges from where they spread out in different directions. Intrusions into the plains of this country possibly occurred from other places, for example the Great Caucasus range. On December 6, 1930, a dead female was found near Kharmandal on the Mugana (Sokolov, 1931). Even earlier, at the end of the last century, leopards were caught in the steppes near the city of Salyanyi (Radde, 1899). On February 20, 1946, a male was killed on the northern Apsheron Peninsula near Bil’gya, 25 km north of the city of Baku. During a severe snowstorm the animal entered a densely populated region and fell into a dry well (Dzhafarov, 1946; Alekperov, 1947). During the last century and in the 1950’s leopards have been caught near Tbilisi (A.A. Sludskii). Leopards used to cross into Turkmenia from Iran and Afghanistan but not regularly and not at a given time of year. Information about regular > = Fig. 121. Leopard track in mud. Cartridge placed for scale. Badkhyz preserve, southern Turkmenia. June, 1961. Photograph by Yu.K. Gorelov. 201 203 260 migrations into Afghanistan (Flerov, 1932) has not been substantiated. In this area animals from the Kopet-Dag foothills sometimes entered the plains but usually traveled only a few kilometers (V.G. Heptner). In the last century leopards were seen occasionally in southern and southwestern Trans-Baikal (Cherkasov, 1884; Baikov, 1927). One was killed within Trans-Baikal in 1912 (Baikov, 1927). By the 1950’s their population has evidently increased somewhat. At the end of March, 1952, at Nizhnyaya Bareya of the Nerchinsk-Zavod region of Chita district, two attacked a flock of sheep and one was killed. Leopards intrude into Trans-Baikal from the Great Khingan range of northeastern China. Leopards were occasional visitors on the midreaches of the Amur (Przheval’skii, 1870), on the Kumara River (Khumaerkhe), on the Amur- Zeya plateau at 51°45’ М. lat. (Baikov, 1927), along the Tyrma River (left tributary of Bureya (Middendorff, 1867), and in the Bureinskii range (Radde, 1862). Very exceptionally one was sighted in November, 1950 on the middle course and at the mouth of the Bira (left tributary of the middle course of the Amur in Birobidzhan), Leopards are not known to people living in the Soviet Little Khingan range, which would indicate that this animal rarely intrudes into this region (Rakov, 1965). ия Fig. 122. Leopard track near spring in Nardevanly settlement. Size of track 12 (width) cm x 11 cm. Badkhyz preserve. Turkmenia. April, 1961. Photograph by Yu.K. Gorelov. 261 In the basin of the lower course of the Ussuri leopards formerly penetrated to Khor (around 48° N. lat.) (Baikov, 1927). In recent decades none has been seen that far north and reports of more southern visits in this region are rare. In November, 1954, one animal was sighted in the upper reaches of the Pervoe Sed’moe (right tributary of lower Ussuri) about 30 km south of Vyazem. In October, 1939, a lone leopard was killed in the Levyi Podkhorenok basin and in October, 1954, another sighted in the middle reaches of the Kanikheza (tributary of the middle Bikin) (Rakov, 1965). In the basin of Lake Khanka, at Beichikhe, leopard intrusions were recorded in January, 1957, and December, 1960. These predators usually penetrate there from China on the trail of migrating roe deer (Rakov, 1965). It has been suggested that most of the leopards sighted in the Primor’e and other regions of the Soviet Far East were intruders from the Korean Peninsula and China and did not breed within the confines of the Soviet Union. To date no litters of these cats have been reported in the USSR (G. F. Bromlei). In Suputin preserve they are sometimes seen in summer, arriving trom the south, but remain for a very short period before returning to their usual habitat (Bromlei and Gutnikova, 1955). On the other hand, leopards not only live regularly in “‘Kedrovaya Pad’ ’’ preserve (south- western Vladivostok) but also breed there (А.С. Pankrat’ev)- Reproduction. Very little is known about this process among leopards under natural conditions, but it has been studied well in captive animals. In the Caucasus these animals usually become sexually aroused in January (Vereshchagin, 1942), in Turkmenia, in the Kopet-Dag, in December-January and again а! the end of November (V.G. Heptner), and in Primor’e (‘““Kedrovaya Pad’ ’’ preserve) (Vasil’ev et al., 1965) and northeastern China (Manchuria) in January (Baikov, 1914). Judging from the dates of sightings of small cubs, mating in the USSR may also occur in other seasons of the year. A few males may follow a single female in estrus and serious combats, accompanied by high-pitched cries, ensue among them. In Kopet-Dag groups of up to five animals have been seen (V.G. Heptner). A female in heat urinates frequently, rolis on her back, arches her back in a characteristic manner whereby the rump is raised, makes typical rumbling and purring sounds, and stops feeding temporarily or eats very little. Among animals held in zoological gardens estrus has been detected at various times of the year, and at intervals of every two to three months if the female happened to remain barren (sometimes it occurs each month, or only twice a year). Estrus lasts 12 to 18 days and in exceptional cases up to 25 days. When the female is stimulated the animals mate many times a day and the intervals between consecutive copulations may be three to four minutes to a few hours (Shereshevskii, 1940). Gestation requires 90 to 105 days, but usually 92 to 95 (Shortridge, 1934; Shereshevskii, 1940). 204 Fig. 123. Trail of а leopard (female) which had been frightened away from her prey. “‘Kedrovaya Pad’ ’’ preserve, near Vladivostok. April, 1965. Photograph by A.G. Pankrat’ev. Parturition occurs mostly at night and the process normally requires six to ten hours. Everywhere within the USSR and also in northeastern China cubs are seen most often at the end of March to April and in April and May (Primor’e, ‘*Kedrovaya Pad’ ’’ preserve; Vasil’ev et al., 1965) but in Kopet-Dag usually in May (V.G. Heptner). Sometimes cubs are found in other months however. Thus, on December 11, 1953, in Badkhyz preserve, in the Gyaz’-Gyadyk mountains (southern Turkmenia) in Nerdevanly gorge, two cubs with their eyes still closed were found, one of which opened its eyes only on December 15 (Rustamov and Shcherbina, 1957; Yu.K. Gorelov). Evidently in the 204 Fig. 124. Individual track of a leopard (female) on fine, wet snow. Match box for scale. ‘‘Кедгоуауа Pad’ *’ preserve. April, 1965. Photograph by А.С. Pankrat’ev. southern part of the leopard range, no definite season for parturition exists. In northern China cubs are usually seen in spring while south of Hwang Ho littering may occur at any time of the year. In Yunnan province cubs are more often sighted in spring but young have been reported in other seasons also. In India cubs are generally born in February or March (Blenford, 1888—1891*). In zoological gardens cubs appear at any time of year. In London Zoological Garden they appear more often at the end of February to early March, followed by end of April to May, June and July, end of September to October to the first half of November, and from the second half of December (Tsukerman, 1953**). The number of cubs in a litter in the southern part of the range and in zoological gardens varies from one to four, and in exceptional cases reaches six. In China a litter consists of one to five cubs, in the northern provinces more often three or four, and in the southern (Yunnan) usually two. In the wilds a lone cub is encountered very rarely. In eastern Asia a litter usually consists of two cubs (Tate, 1947). The number of young in a litter within the Soviet Union has not been established for want of adequate . data, but most often consists of two or three. In the western Caucasus a *(Sic]; Blanford. Not in Literature Cited—Sci. Ed. **(Sic]; Zuckerman. Not in Literature Cited—Sci. Ed. 205 264 female with two cubs was caught (Dinnik, 1914). In the Geok-Tepi region of Turkmenia a female with two young was sighted. In Badkhyz two litters with two cubs each were found (Filippov, 1936). In February, 1959, near Ashkhabad a female with a single cub was caught (Polozov, 1959) and in the same region in December another with two cubs trailing behind her. Families consisting of three members have been sighted time and again in Turkmenia (Dement’ev and Rustamov, 1956). The number of cubs there is usually two but may reach four (V.G. Heptner). In Primor’e, in ““Кеагоуауа Pad’ ’’, litters comprise one to three cubs (Vasil’ev et al., 1965). Growth, development, and molt. Cubs are born blind and helpless but covered with dense and fairly long hair. The main color tone of the coat in the upper part of the body in newborns from the nose to the tip of the Fig. 125. Tracks of a large leopard (male) on wind-hardened and water-logged ^ snow; snow has drifted into the tracks. Kedrovaya River in ‘‘Кедгоуауа Pad’ ’’ preserve, near Vladivostok. January, 1960. Photograph by A.G. Pankrat’ev. 205 we) n N Fig. 126. Sketch of foot pads on sole of front (left) and hind paws of a leopard killed on Cape Gamov at Vladivostok. 1930. Sketch by G.D. Dul’keit. tail is light brown. This background is uniformly speckled with tiny dark brown and black spots which do not form rosettes (Fig. 127). With age these spots fuse into rosettes. They are larger on the back than on the flanks. The underside of the body and the tail are creamy-white and speckled with large blackish-brown spots. The weight of a newborn cub is 500 to 700 g and body length about 15 cm. The young open their eyes on the 7th to the 9th day. On the 12th to the 15th day they begin to crawl around the den and by the second month emerge from it to play nearby. At this age they also begin to eat meat and shed the juvenile coat. Initially they are fed on milk alone, then semidigested meat regurgitated by the mother, and finally birds and small mammals. After one year the young still differ greatly in appearance from adults; in February (1959) the length of the trunk of a dead female was 117 cm and that of her cub was only 73 cm (Polozov, 1959). The cry of a leopard cub is very similar to the meowing of domestic cats. Lactation continues for five or six months. As soon as the young begin to eat meat the mother takes them with her to hunt. Cubs sometimes accompany their mother until she comes into estrus. Young leopards, deserted by their mother, continue to live together and follow adult animals, consuming the remains of their prey. Hence groups of two to five are sometimes sighted. At the end of winter juveniles separate from each other and usually abandon the region in which they were born. The male lives with the female only during the period of rut. References in literature about the male remaining with his offspring are erroneous. Sexual maturity sets in at the age of 2 to 3 years and ability to reproduce 207 205 Fig. 127. Cub (male) of an Amur leopard. Sanduga River, west shore of Amur Gulf near Vladivostok. February 27, 1956. Photograph by А.С. Pankrat’ev. continues up to 12 to 15 years of age. In captivity some animals have survived for 21 years. In the London Zoological Garden 52 leopards lived on the average of 39.3 months each, but one animal survived for 13 years 8 months (Mitchell, 1911). Molt in leopards has not been studied. Enemies, diseases, parasites, mortality, and competitors. Leopards have no enemies except tiger, and gray and red wolves [dhole]. In India red wolves sometimes gather in packs of 30 to attack a leopard, which tries to rescue itself by hiding in trees (Anderson, 1964). Diseases suffered by wild leopards have not been studied. Only one instance of a rabid leopard is known in the Caucasus. In zoological gardens this predator suffers from all the diseases characteristic of other feline species (infectious enteritis, carnivore distemper, etc.). Three species of helminths are known to parasitize leopards in Azerbaidzhan: Taenia hydatigera, Mesocestoides lineatus, and Ancyclostoma canium (Asadov, 1947; Sadykhov, 1955). These parasites have been reported additionally: Toxocara mystax (Mozgovoi, 1953), Dirofilaria granulosa (Skryabin and Shikhobalova, 1948), and roundworm Dracunculus persarum, a subcutaneous parasite of man (Pavlovskii, 1935). Tiger, gray and red wolves, and possibly striped hyaena, represent serious competitors of leopards. In Primor’e it has been noted that at places where 206 267 tigers reside, leopards are absent ог very rarely encountered. In India instances of leopards being killed by tigers have been recorded. It is known that hyaenas chase leopards away from their prey (Pocock, 1939). Once:a leopard was mortally wounded by a porcupine (Burl’er, 1955)*. Population dynamics. This has not been studied. It is only known that due to killing by man and a reduction in its food base (destruction of ungulates), the leopard population decreased considerably during the last century. In the Caucasus, Tadzhikistan, Uzbekistan, and the Far East this species is on the verge of total extinction. Field characteristics. The leopard is a large cat weighing 30 to 75 kg. It differs from the ounce in its yellow or reddish-yellow background coat color, speckled with small, bright, black spots disposed in rosettes over most of the body. Its tail is equal to two-thirds of the body length. Its tracks are smaller than those of a tiger (Size of a track on snow 12 cm x 12 cm) but far larger and rounder than those of a cheetah. Claw impressions, unlike cheetah, are not seen in leopard tracks. Tracks extend in a uniform chain with a distance of 40 cm between individual marks (Figs. 121 to 125). A leopard conceals the remains of its prey under the roots of fallen trees, in patches of wind-felled trees, in upright trees, in hollows, and in rock crevices, and feeds on it for two to four days. In Primor’e the head is invariably absent from all prey items, even when the carcass has not been fully consumed (Vasil’ev et al., 1965). The intestine of victims along with their contents are eaten, and even highly decomposed carcasses —which is not the case among other species of cats. Leopards bury their feces. Leopards are fairly silent animals and their call heard only rarely. During the breeding period they call almost every night. At other times their call can be heard in the mountains over a distance of several kilometers. The leopard’s call consists of short, low-pitched ‘‘zzugghs’’ similar to the sound of a saw, delivered at short intervals; sometimes the calls are prolonged and almost merge with one another. During the breeding period the call resembles a harsh cough which turns into labored breathing. On approaching a water hole leopards often purr. When disturbed they rumble and snort. On falling into a trap the animal maintains silence. (A.S.) Practical Significance Leopards are fur-bearing animals but their skins are not handled in the fur trade of the Soviet Union because the population of this species is too small. At the end of the last century in one of the former Krasnovodsk districts (western Turkmenia) only 10 leopards were caught annually (Silant’ev, *[Sic]; Bourliere. Not in Literature Cited—Sci. Ed. 268 1898). In Turkmenia the following numbers of skins have been prepared: 4 in 1925-1926, 5 in 1926-1927, 9 in 1927-1928, and 7 in 1928-1929 (Kogan, 1931). Later the catch in Turkmenia increased: 6 skins were tanned in 1948, 7 in 1949, 5 in 1950, 10 in 1951, 6 in 1952, 9 in 1953, 8 in 1954, 9 in 1955, 41 in 1956, 24 in 1957, 35 in 1958, 19 in 1959." In 1912 only 2 leopards were killed in the Amur region, 11 in Primor’e, 1 in Trans- Baikal, and about 15 in northeastern China (Baikov, 1927a). Only 2 or 3 were caught some 10 to 12 years later in the Ussuri territory (Solov’ev, 1925). According to current data (1960’s) published by the fur industry the skins of this species are not listed separately but grouped with ounce and cheetah under the general classification of “Чеорага””. In the 1930’s some 120 ‘Чеорага”” skins were tanned annually (Kogan, 1931), most of which were ounce, and only 20 leopard. The total output of leopard skins per se for the Soviet Union up to the 1930’s was placed at 20 to 30 pieces per annum (Kuznetsov, 1932). In recent decades 20 to 30 skins have been tanned _ in a season but some retained by the people. In the Leningrad fur auction of 1956, 40 leopard skins were put up for sale and sold at an average price of $10 each (American currency) (Kaplin, 1960). The contribution of the U.S.S.R. to the world trade in leopard skins is obviously insignificant. The annual world output from 1907 to 1910 was 10,000 to 11,000 skins, of which 5,000 came from Asia and the rest from Africa (Brass, 1925). In the 1920’s the annual world output dropped to between 1,000 and 2,000 (Zhitkov, 1928). In the 1950’s leopard skin coats and other apparel became fashionable and the world output in these years soared to 20,000 skins (Shou, 1958). The fur market in Kunming alone (Yunnan) handled 4,000 to 5,000 skins annually (1958; A.A. Sludskii). A leopard’s coat versus one of other skins, reckoned at 100% durability, is quite durable with a rating of 75% (opposed to fox with a rating of only 40%). Different parts of a leopard’s body are widely used in Chinese medicine even today and hence, apart from skins, even the bones are marketed in China. Limb bones are the most valuable. Desiccated eyes are also highly prized. In fact, a hunter often realizes more from the sale of leopard bones and eyes than he does from the sale of its skin. In the Soviet Union the damage inflicted by leopards on game has been very small because the population of this predator has never been large. Only recently have leopards been classified as a dangerous enemy of cattle in Turkmenia and in the Ussuri territory, but the damage caused by this predator is negligible. In the past this cat threatened game even in the Caucasus and 10 is possible that some cheetah skins were included in the leopard count in Turkmenia. One must also remember that a large number of leopard skins are retained by hunters. Up to 15 leopards were killed annually in Kopet-Dag alone in 1940 and 1941 (V.G. Heptner). 208 269 Trans-Caucasus. Compiaints about losses were registered in Lenkoran as late as January, 1930 (Starichikhin, 1930). In the Far East leopards were formerly responsible for losses among sika deer in state farms. Among the enemies of sika husbandry, especially in Pos’et region, leopard ranked second, with wolf coming first. Reportedly, a leopard once sneaked into a deer park by crawling through a wire net fence 3.0 meters thick, consisting of 16 rows of barbed wire (Menard, 1930). In India, where these predators are still quite numerous at places, attacks on humans have been reported. In fact some animals have become man- eaters to such an extent that they have threatened the safety of human beings over a wide area. By the end of 1850’s one leopard had killed about 200 people in a three-year span (Pocock, 1939). In Kumaon in the present century two leopard man-eaters killed 525 people before they were put down (Corbett, 1957). A leopard becomes a man-eater by accident—when the destruction of game by man deprives it of normal food. Moreover, unlike tiger, leopards will sometimes eat carcasses. At certain places in India dead bodies are abandoned during epidemics; the animal coming across such human corpses feeds on them and becomes accustomed to human flesh. When the epidemic had ended, it continues to attack living persons. Old and crippled animals, who cannot hunt ungulates successfully, also turn into man-eaters. Man-eaters have also been reported in China but are extremely rare (Allen, 1938). In the Soviet Union neither biological nor social conditions have given rise to man-eaters either in the past or present. Usually only an injured leopard will attack man (very few cases); nevertheless instances of an uninjured normal animal attacking man have been recorded. In the Caucasus during the winter of 1881/1882 in the Groznyi region a 1еораг4 tore a man to bits in the forest. In 1875 near Sochi one attacked a sleeping worker, severely bit his occiput, and attempted to carry him away into the forest, but fortunately was repulsed by other workers (Radde, 1899). In the 1890’s in the Lenkoran lowland near Prishib village another leopard killed a 15-year-old shepherd boy (Dinnik, 1914). In Turkmenia two incidents have taken place. Early in the present century in a gorge in western Kopet-Dag, a leopard unexpectedly ambushed a man from behind a clump of rushes. In the life-and-death struggle which ensued the man managed to strangle and stab his attacker. It was later discovered that the leopard was old with worn-out canines. The hunter was further saved from serious injuries partly because he had on a heavy coat [khalat]. In the early 1920’s a hunter from Verkhne-Skobelev village in hot pursuit of rock partridge in the Kopet-Dag jumped onto a stone behind which a large leopard happened to be sleeping. The animal sprang instantly to the attack and, placing its forepaws on the hunter’s shoulders, tried to seize him by the 208 270 throat. Luckily, the gun with the hammer cocked, which the hunter was carrying in his hands, jammed in the attacker’s throat, and the predator died within a few seconds. In my opinion (V.G. Heptner) this incident should not be viewed as a leopard attack. By and large, leopards are far less dangerous to man in the Soviet Union than wolf; this predator has killed quite a number of people in this country (see Volume II, Part 1 of the present monograph. In the Kopet-Dag, a leopard has sometimes tracked a man as though pursuing him, but has not attacked (V.G. Heptner). In the Caucasus leopards were shot on chance encounters or caught in traps set up mainly in trees standing in water or on specially made bridges. Strychnine administered in the form of pellets embedded in the remains of prey was also used. In Turkmenia leopards were shot and caught in traps Fig. 128. Catching of live leopards in the Far East. Hunter nudging one taking refuge in a tree with a pole (from Sysoev, 1955). 209 209 OA set up on trails. In the Far East native people were as afraid of leopard as of tiger. Only rarely was one caught by setting up bows and guns on trails and at the site of remains of a kill. The animal was also shot in chance encounters, caught in traps, poisoned with strychnine, and chased on fresh snow with the help of dogs. In the 1920’s some deer farmers used German sheepdogs for hunting leopards in deer parks and near them. These dogs wore spiked neck collars to protect them from seizure by the predator. Two or three dogs spotted an animal and chased it up a tree where it was then shot. In the hunting season of 1927 one hunter (A.A. Bogoyavlenskii) with two dogs caught three leopards (V.V., 1927; Vereshchagin, 1929). A leopard quickly kills pursuing dogs by biting their skulls. The predator is highly resistant to wounds and it is a rare feat indeed when one is felled with a single shot. Fig. 129. Leopard nudged down from a tree and pinned to the ground by hunters (from Sysoev, 1955). In the All-Union standards on skins in the 1940’s to the 1960’s leopard skins were classified according to size, grade, and flaws. The raw skin of a leopard in the last decade was not prized and, depending on grade, 5014 for 80 kopecks to 4 rubles 70 kopecks. Formerly, it was more expensive; for example, in the 1860’s in the Ussuri territory a good skin sold for 15 to 20 silver rubles (Przheval’skii, 1870). Skins were used in making rugs and sleigh robes. Apart from skins, in the Far East the Chinese formerly valued other body compcnents (meat, bones, blood, brain, bile, teeth, and claws). A male was priced higher than a female and an adult higher than a cub. The average price of a freshly frozen carcass was two to three gold rubles per kilogram. Different parts of the carcass were used in Chinese medicine and as talismans. Leopards, like tigers, are especially honored by natives of the Far East and always worshipped. Spotted idols used to be set up on altars in high altitude passes (Baikov, 1927). Udekheits and Orokhets of olden times idolized this predator and its 14015 were found among them (Schrenk, 1858; Przheval’skii, 1870). In recent decades, due to the great demand for live leopards, they have been trapped or caught live without traps. In the Ussuri territory hunters catching tigers (see section ‘‘Tiger’’) also caught leopards. On finding fresh tracks of a leopard, hunters set their dogs in pursuit. When the dogs have stopped the animal or chased it into a hiding place, hunters pin it to the ground with forked poles and then truss it (Sysoev, 1955). Ifa leopard escapes the dogs and climbs onto a tree, the hunter nudges it down with a pole and his companions pin it to the ground. Not only young leopards but also adults are caught with forked poles. Leopards survive and breed well in captivity. in the Soviet Union this big and beautiful predator, like the tiger, is an asset to the mountains and forests in which it lives. At present it causes no harm to cattle or game. It also poses no danger for man if not attacked. Leopards should be protected in the Soviet Union since this species is of great scientific and aesthetic value. To protect the leopard and prevent its total extinction, hunting and catching live animais should be banned throughout the Soviet Union.!!? International conventions for the conservation of leopards should be established with neighboring governments (Korean Democratic Republic, China, Afghanistan, Iran, and Turkey). Reserves and preserves ought to be ‘11Such a measure had already partly arrived in Primor’e territory in 1956 and shooting of leopards banned year-round in Khasansk, Vladivostok, Shkctov, Budenov, and Lazov regions, and within Vladivostok, Artem, Suchansk, and Nakhodka municipalities. A penalty of 100 rubles is now imposed for catching a leopard. Contrarily, in other republics (Armenia, Azerbaidzhan, Turkmenia, and Tadzhikistan) killing leopards continues year-round and a prize is even awarded for any animal killed. 21 set up for the conservation not only of this predator, but also of the animals on which it survives (mountain sheep, roe deer, sika deer, etc.). Raising leopards should be organized in zoological gardens by creating pedigree groups of a single subspecies. In Africa this predator has almost totally disappeared over large expanses in recent years because leopard skin coats became fashionable in the 1950’s and 1960’s. Subsequently wild pig and baboon, the main food items of leopards, multiplied unchecked; the loss inflicted on crops by these animals far exceeded any loss to domestic animals caused by leopards in the past. For this reason, and under intensive pressure from campaigning ecologists in several African states, the leopard was removed from the category of dangerous animals and placed on the list of animals to be protected by law (Arrua, 1958*). In some countries (Kenya) work is now in progress to reacclimatize the leopard (Right, 1960**). (A.S.) *(Sic]; not in Literature Cited—Sci. Ed. **1 Sic]; Wright? Not in Literature Cited—Sci. Ed. Genus of Snow Leopard, or Ounce [Irbis] Genus Uncia Gray, 1854 1854. Uncia. Gray. Ann. Mag. Nat. Hist., 14, p. 394. Felis uncia Schreber, FD: Large-sized animals. Legs not long, trunk very long, general build stocky, sacral region high, higher than shoulders, and profile of back slopes forward somewhat (appearance typical of small cats, Felis; see Fig. 30). Tail length distinctly more than half length of body. Claws wholly retractile and paw structure same as in big cats (Panthera, see Fig. 278). Body covered with dark annular spots. Skull quite powerful with tuberosities, crests, and powerful zygomatic arches, but less massive, less heavy, and with less developed crests than genus Panthera. Skull broad and short with large orbits set high; interorbital region distinctly elevated, with upper line of profile sharply curved, falling relatively steeply fore and aft of interorbital region. In general, skull bulges notably. Facial region shorter than in big cats (genus Panthera); distinct notch occurs at places where nasals join frontals. Nasals short and broad; their anterior width almost equal to their length along midline. Basicranial axis markedly raised toward the front (not horizontal), meets basifacial axis (axis of facial portion of skull) and forms a distinct angle. Postorbital constriction distinct and broad (broader than interorbital). Cranium relatively more voluminous and enlarged than in big cats (genus Panthera), not elongated, or slightly so, with facial region of skull relatively less powerful. Orbits open. By and large skull resembles that of small cats (genus Felis). Tympanic bulla relatively small and slightly expanded. Distance between it and posterior wall of glenoid fossa small; the two almost touch each other. Antero-outer (ectotympanic) chamber of bulla very well developed; two chambers nearly identical in size, with suture between them shifted away from auditory meatus (Fig. 23). Anterior part of entotympanic chamber very narrow in front. Deep depressions present on basioccipital portion of occipital bone in front of jugular foramen on each side, internal to tympanic bulla. Structure of hyoid apparatus same as in big cats (genus Panthera), i.e., the epihyal in the form of an elastic ligament (Fig. 31). Chin portion of lower jaw set steeply (almost at a right angle). Dentition complete (second premolar present); teeth, especially canincs, relatively less massive than in big cats (leopard). 212 2S Sexual dimorphism absent. This predator generally catches prey as large or even much larger than itself. It hunts by lying in wait. It is a solitary animal and lives in mountains, mainly in high altitudes up to snowline. In spite of the structure of the hyoid apparatus, similar to that of big cats (genus Panthera), the *‘roar-growl’’ type of call is not heard. It “‘purrs’’ during both inhalation and exhalation (as in small cats, genus Felis). It tears its prey in the manner of big cats but its eating posture is that of small cats (genus Felis, see Fig. 32). The range of the genus (see description of the species) is relatively small, somewhat less than that of other genera. It covers montane Central Asia as far as the Himalayas in the south, in the west to the Hindu Kush, Pamir-Gissar system, and Tien Shan, in the north to the Altai (probably to [lake] Baikal in the past), and in the east includes eastern Tibet. The genus Uncia is unique in several respects and placing the ounce in genus Panthera is not justified. Craniologically it differs from species of the genus Panthera more than the latter differ among themselves in any combination. At the same time, genus Uncia shows a combination of several morphological and ethological characteristics of both big cats (genus Panthera) and small cats (genus Felis). Thus the structure of the cranial region of the skull is extremely similar to that of large species of the genus Felis, but a hyoid typical of genus Panthera, eic. Ethologically, as far as can be judged from observations of captive animals, the ounce is closer to small cats (genus Felis) and essentially identical with their ‘‘peripheral’”’ species —the clouded leopard (F. nebulosa; Hemmer, 1964 and 1966). This species has also been placed time and again in “‘subfamily Pantherinae,”’ but it is now clear that this is a genuine small cat although it shows some specialized features. It is sometimes placed in a separate genus (Neofelis) even by authors who recognize only a few genera in the family (Ellerman and Morrison-Scott, 1951 and 1966). Features of the ounce, discussed above as an intermediate form between genera Panthera and Felis $. 1., define its taxonomic position and, simultaneously, highlight the difficul- ties of a clear separation of genera of present-day cats (excluding the cheetah). Paleontological data pertaining to the origin of the genus are not avai- lable. Quite possibly its origin, like that of the genus Panthera, should be placed at the end of the Pliocene or in the Pleistocene. There is only one species in the genus, ounce or snow leopard, Uncia uncia Schreber, 1775, representing 3.5% of the species of the family. The ounce is a valuable game and fur-bearing species and one of the prized possessions of zoological gardens. It exerts a control over the population of some mountain ungulates. In the USSR the one species in the genus constitutes 9.0% of all species 276 of the family inhabiting the Soviet Union and 0.3% of mammals of our fauna. The geographic range in the Soviet Union covers the montane regions of the eastern parts of Middle Asia and part of the extreme south of middle Siberia. This species is an object of sport and also important to the fur industry. It is mainly valued, however, as an object of trade in live animals for zoological gardens. The damage inflicted by the ounce on livestock is very little and can be ignored. (V.H.) SNOW LEOPARD, OUNCE [/RBIS]! Uncia uncia Schreber, 1775 1775. Felis uncia Schreber. Saugeth., tab. 100, vol. 3, р. 586 (1778). Type locality not indicated.? 1830. Felis irbis. Ehrenberg. Ann. Sc. Nat., vol. 21, pp. 394 and 406. Substitute for F. uncia Schreber. Altai. 1855. Felis uncioides. Horsfield. Ann. Mag. Nat. Hist., vol. 16, p. 105. Nepal. 1950. Panthera uncia schneideri. Zukowsky. Zool. Gart., vol. 17, p. 213. Sikkim. (V.H.) Diagnosis Uncia uncia is the only species of the genus. Description Size large, roughly corresponds to that of leopard (Panthera pardus), but slightly smaller on the average. Trunk very long and stocky; general appearance similar to that of small cats (genus Felis, Fig. 30), 1.е., with a slightly raised sacral region. While generally resembling the leopard, the ounce also differs in its less strongly ‘For a correct Russian name for this species, see footnote по. 1 under “‘leopard’’. From a scientific viewpoint it is incorrect to call this animal simply ‘‘leopard”’ [bars]. 2S I. Ognev (1935; followed by Stroganov, 1962), based on some indirect evidence, suggested that Iran should be regarded as the type locality. His conclusions are not adequately substantiated; in particular the occurrence of ounce in Iran has not been generally proved; evidently this animal is absent there or is only found as an exceptional rarity (vagrant—?, see *‘Geographic Distribution’’). In any case, Iran as the type locality is inadmissible. The far better substantiated suggestion of R. Pocock (1930), to consider the Altai the type locality not only for the Soviet Union but also Mongolia [sic], should be accepted. (V.H.) 277 developed anterior (chest) portion of the body and a relatively small and more rounded head. Upper line of profile of head bulges, eyes quite large, and pupils round. Ears short with rounded tips, broad at base, almost hidden within the coat in winter; long hair (tuft) at tips of ears absent. Mane or long hair on cheeks (whiskers) also absent (Fig. 130). Vibrissae white and black and up to 105 mm long. Tail very long, relatively longer than that of other Soviet cats, and more than three-fourths length of body. Paws broad and massive; claws a light horn color. In spite of having dense and long hair, the animal with its somewhat flattened and less muscular trunk appears less massive than the leopard (Figs. 132 and 133). Forepaws larger and more powerful. ‘The winter coat is very rich, soft, and silky. Tail covered with very long and dense, lax hairs, and appears very thick and fluffy. Pelage of the ounce, compared to that of other species, moderately dense, long, and soft. On the back, density about 4,000 hairs per cm?, with eight underfur hair to one body hair. A characteristic feature is the absence of any significant difference in the length of different categories of hair; guard hair only 1.2 times and body hair of category Г just 1.1 times as long as hair of underfur. Body hair of category IV and especially that of category Ш somewhat longer than body hair of categories I and II. Length and thickness of guard hair on the back 54 mm and 77 microns respectively; length of body hair (categories I to ТУ) 48, 47, 51, and 49 mm and thickness 76, 58, 43, and 34 microns; length of underfur hair 43 mm and thickness 20 microns. Type of fur abdominal’’* (B.F. Tserevitinov). In pelage density the ounce differs from all other big cats and is more similar to small cats (genus Felis), occupying a somewhat intermediate position between them. Its being ‘“‘intermediate’’ is important in any comparison with small cats. These differences, however, are hardly important in taxonomy and are more associated with conditions of the habitat of the animal and its thermal requirements (thermoregulation). The contrast between winter and summer coats is very great, far more than the corresponding contrast in the leopard. General color of the main background of winter coat very light gray, almost white, or whitish with a smoky tinge, more perceptible on back and up the flanks; it may also be light yellowish. Lower part of flanks, whole of underside of body, and inner surface of legs even lighter, being almost white or with a yellowish tinge (Fig. 132).3 *Initial quotation mark for this quote missing in original Russian; meaning of last sentence unclear.—Sci. Ed. УГве yellowish shade seen in skins of some animals which died in captivity (Zoological Museum, Moscow University) is possibly an artifact. РЫ Fig. 131. Skull of snow leopard, Uncia uncia Schreb., №. $ 85724. Collection of the Zoological Museum, Moscow University. Trans-Ili Alatau, Tien Shan. August 4, 1969. Sketch by N.N. Kondakov. There are no distinctly yellow or reddish or tan shades in the color 215 scheme of the ounce. On the main background are scattered dark spots, some of which are annular, with a diameter of up to 7.0 to 8.0 cm, and others large or small and solid (Fig. 133). Spots are sparse, far more sparse than in the leopard. 280 Fig. 132. Snow leopard. Vil’nyus Zoological Garden. Photograph by В.В. Marm. Solid spots of different sizes are scattered on the head (smallest) and neck and legs (larger ones above, smallest below) where annular (rosette) spots are absent. The latter are rounded or somewhat elongate and disposed on the back, flanks, upper portion of the thighs, along the shoulders, and on top of the proximal portion of the tail (Sometimes in two rows here). On the posterior part of the back, spots often fuse into short transverse stripes. Between the annular spots a few small solid spots are scattered. Large solid spots on the distal half of the tail often surround it in the form of an incomplete ring set in a transverse direction; the very tip of the tail is usually black. The color of the central areas of annular spots corresponds to the main background in a given section of the body. The spots are black but their outlines are not sharp in a dense coat, and are often greatly blurred; their color is therefore not complete and appears more as a dark gray shading. This is particularly true of the annular spots. Solid spots, especially the large ones, are more sharply outlined and a more intense black color. In the summer coat the general background appears more light-colored, almost white, and the outlines of the dark spots sharper, darker, and usually pure 217 Fig. 133. Snow leopard. Vil’nyus Zoological Garden. Photograph by B.B. Marm. black. The smoky film is fainter than in the winter coat.* The pattern of spotting shows some individual variability in density of spots, etc. The color of the spots may also vary (Figs. 134 and 135). Usually they are pure black but sometimes brownish. The main background color of the coat is less variable. In general, however, individual color variability in both these features (color and pattern) is comparatively less marked in the ounce than in the leopard. Sexual dimorphism in color is absent. The young in their first coat (newborn) differ sharply in the depth of pigmentation of spots, which are few in number, and also in the presence of few annular spots. Large black or brownish solid spots are present on the ‘The characteristics of the summer pelage of the ounce and its variability is not adequately known. The animal carries it for a very short time and most pelts in museums are of winter skins. Usually, therefore, winter skins are described. 282 Fig. 134. Variation in color in winter coat of snow leopard, Uncia uncia Schreb. 1—No. S 35130, female, February 1, 1939. Dzhety-oguz, 55 km south of Przheval’sk, Tersk-Alatau, western Tien Shan; 2— No. S 35128, male (not adult), February 1, 1939, same places as above; — Мо. S 35126, female, February 1, 1939, Dzhetyoguz; 4—No. S 72736, male, December 19, 1964, western Tien Shan; 5 — №. S 72958, female, March 20, 1965, western Tien Shan; 6— No. $ 35127, female, February 1, 1939, Akshiprak area, 90 km southwest of Przheval’sk, Tersk- Alatau, western Tien Shan. All skins from collection of the Zoological Museum, Moscow University. Photograph by V.G. Heptner. 219 Fig. 135. Variation in color in summer coat of snow leopard, Uncia uncia Schreb. 1—No. 49243, August 13, 1939, Tien Shan; 2—No. 49240, female, June 13, 1940; Tien Shan; 3 — No. 49953, female, 1945, Mongolia; 4— Мо. 49242, male, August 8, 1939, Tien Shan. All skins from collection of the Zoological Museum, Moscow University. Photograph by V.G. Heptner. 284 back and similar short, longitudinal stripes in the rear (see Fig. 150). Differences from the leopard are quite distinct. By the first winter the coat of cubs does not differ from that of adults.° Geographic variation in color is absent. Apart from the features discussed above (see characteristics of the genus), the skull of the ounce differs from that of leopard, with which it bears features of similarity, in the following respects. Skull relatively broad and short with widely separated zygomatic arches; cranium more enlarged and roomy with a relatively broad postorbital region (its width about 30% of condylobasal length and usually more than interorbital width), reflecting affinity with small cats (genus Felis) and not big ones (genus Panthera); and skull wider in interorbital region. The above results in a more rounded skull which, on the whole, is similar to that of small cats. Orbits placed somewhat higher, and more rounded; zygomatic process of squamosal stops far short of base of postorbital process of jugal; lower anterior edge of orbits not thickened; hard palate shorter and broader; rostrum broader but its width above canines not greater than postorbital width; and choanal edge of palatines with two small pointed projections along sides of suture between palatines. Anteriorly, nasal process more rounded; interpterygoid vacuity wider; paroccipital processes thin and long, project markedly beyond surface of tympanic bullae, and tips pointed and bent forward in form of a hook. Jugular foramen relatively small and rounded. Measurements for adult animals (males and females combined): body length (10) 103 to 125 cm (M 112.1); tail length (10) 80 to 105 cm (M 91.8); length of hind foot (8) 22.5 to 26.0 cm (M 23.8); and length of ears (8) 6.6 to 7.2 cm (M 7.0). Weight (5) 22.0 to 39.0 kg (M 33.0) (material from the Zoological Museum, Moscow University, and data of A.A. Sludskii and B.B. Marm). Female (Talassk Alatau, April): body length 125 cm, tail length 90 cm, hind foot length 24.5 cm, ear length 6.6 cm, height at withers 50 cm, height at sacrum 60 cm, and weight 39 kg. Maximum body length up to 130 cm and weight up to 40 kg (Stroganov, 1962). Females are evidently somewhat smaller than males. Cubs aged about 9 to 10 months (January and February): body length 85, 82, 87 cm; tail length 75, 65, 76 cm; length of hind foot 22, 21, 22 cm; and ear length 68, 63, 70 cm (first and last from the same litter; Zoological Museum, Moscow University). Skull measurements are given in Table 4. (V.H.) УТве view that the spots of ounce become more diffuse with age (Stroganov, 1962) needs confirmation. It may be true of the first adult coat but is hardly applicable to subsequent coats. (НА) ‘зтечиие 3э1л06 Jo э$04} Wor 1эууир оц ор ($еАвецин) duel э4 418098 oy) Jo syied 1э410 WOIJ DUNO 10} зиэшолазеэш ‘элп]елэ JY) иг вер 1э8вэш шо] 8шарп[ :э10 № 41001 [ет5$еилео 1э44п jo Ч134э] эитиео JNOYIIM MOI 41001 Jo 41349] эииео YIM MOI 41001 JO Ц}8иэ] YIPIM [814101504 Чартл [23192012] рим эцешо8Ай Ч 30э1 |е5еа01Арио) 41319] 1594215) сос ох £981 | 6061 зэешэу 3[Npy зэеш пру (Аутблэли(] №0250 ‘LUMaSNA] [е9180[007, Шол} 5иэциоэ4$) ULYS иэт |, ay) итол} ‘072ип п12и] ‘рле4оз| MOUS JO ззиэшолизеш [[NYS °р ЧЕТ, 555 1эелеЧЭ 5э[ешэу рие soyeyy Systematic Position Uncia uncia is the only species in the genus. Geographic Distribution This species is distributed in the upper and middle zones of mountains in Central Asia, Turkestan, and southern Siberia. Geographic Range in the Soviet Union The range in the Soviet Union (reconstructed) represents the northwestern and northern periphery of the range of the species and covers the montane systems in eastern Middle Asia (Turkestan) and southern Siberia from the Amu-Darya to Baikal. In the west the range of the ounce covers the Pamir-Gissar system and Tien Shan, the entire Pamir, the Darvaz range including its southwestern spurs, the Peter the Great, Trans-Alaisk, and Gissar ranges together with the Baisuntau mountains, the Zeravshan range westward to the districts of Pendzhikent and Turkestansk to the west, at least to the meridian of Zaamin (Zaamin preserve) and the Alai range. Ounces were never present in the Mal’guzar mountains (Fig. 136). The southern boundary fronting on the Amu-Darya in southern Tadzhikistan describes an arc beginning at Pyandzh on the north and encompassing the Kulyabsk, Dashti-Dzhumsk (connecting the Khazrat- Ishi mountains with the Darvaz range), Muminabadsk and Kyzyl-Mazarsk regions, where the animals are regularly met with. From here the boundary runs northwest and encircles Dushanbe from the north. North of that city, the ounce lived along the Varzob. From there the boundary runs along the southern slope of the Gissar range to the west and later southwest along the Baisuntau [range]. In Baisunsk mountains the ounce is distributed in the south at least to the latitude of Deinau on the Surkhan-Darya. This species is evidently absent at Kugitang. In this part of its range (southern Tadzhikistan), the animal occupies the higher portions of the country. It vicariates with the leopard, which occurs here in several places, but to the south of the range boundary described above, and at lower elevations. Farther to the north and northeast, the ounce is met with in all the ranges of the Tien Shan system, to the south including the Kuraminsk and Ferghansk ranges, reaching the Ferghana valley from the north, and westward reaching the western spurs of the Chatkal’sk (Montane-forest preserve near Parkent), Pskemsk, Ugamsk, and Talassk ranges (in the west to Aksu-Dzhebagly preserve). The Karatau, Kirgizsk (Aleksandrovsk), Kungei Alatau and Trans- ‘лэщ4эН “OA “SIOATY ими рие ерП иээлуэа иотЗэл puke ‘еуэО JO ISINOD Jamo] ‘УзлеАоизету “Seq-jadoy — ээцэллиээо $.[ециие dy} 0} $2919191э1 зпоэцонэ э1оцэр $э[ол1 ‘(WY ит э[е9$) UOTUL, JAIAOS ay} и! “49149 вип втри[] ‘ээипо до эвиел Jo Алерипоа илэцузюм “OT ‘8 288 Ш Alatau ranges form the northern limit of occurrence (ounces are encountered on the northern slope south of Alma-Ata). In the Chu-Ili mountains ounces are absent; they inhabit the Dzhungarsk Alatau and its offshoots—the Altynemel’, Katutau and Aktau—and the Tarbatai and Saur. In Altai, the ounce occurs far to the south. In the west they have been met with along the upper and middle course of the Bukhtarma from its uppermost reaches (Beleya Berel’ River and Chindagatui mine) and noticed even as far as the lower Zyryanovsk. They also live on the upper Narym, between Narym and Bukhtarma, and in the Sarymsakty range south of Katon-Katagai (Altaisk station). The more easterly range includes the mountains of the Argut basin (upper Katun), including its tributaries the Dzhassater in the south and Shavla to the north (near the mouth of the Argut on the Katun). The Chuisk steppe and the surrounding ranges, and evident- ly the mountains of the entire Chui basin also fall within the range. Even more eastward the range covers the Bashkaus basin, the left tributary of the Chulyshman, and Chulyshmansk basin in the north, including the Chul’chu and Kygu, its right tributaries. Here (Altai preserve) the range extends to the southern extremity of Lake Telets. Thus, the following main ranges of the southern and, in part central, eastern, and northeastern Altai, and associated massifs fall within the geographic range: Narym, southern Altai, Tabyn-Bogdo-Ola, Listvyaga, Katunsk, Sailyugon, southern and northern Chui; Chikhachev, Aigulansk, and Chulyshmansk mountains; Chulyshmansk uplands; and Shapshal’sk mountain (Chapchal). In the portion of Altai outlined above the distribution of the ounce is uneven—in the far south they are found more or less regularly (upper Bukhtaria, Chuisk mountains), and in the north and northeast, occur as vagrants, very rarely at some places (Lake Telets). Still farther east closely associated with the Altai-Sayan system and Tuva Autonomous District, the distribution includes the Tannu Ol range, region of the upper Kemchik, and the upper Bol’shoi and Malyi Yenisei. Farther north, in the Sayan range, directly adjoining the Shapshal’sk [range] (Altai), the distribution runs to the east at least to the upper Kazyrsuk (right tributary _ of the Yenisei®). It is highly probable that the range included the entire western Sayan system (except the Sayansk, Kurtushibinsk, and Ergak- Torgak-Taiga ranges), and the eastern part of eastern Sayan system— Udinsk range (source region of the Bol’shoi Yenisei). Farther east, the range covers Tunkinsk [range] and possibly the Kitoisk bald (west of the southern end of Baikal) and also Pri-Baikal mountains along °Опе was caught approximately 130 km south of the city of Abakan,.40 km from Веуа (Beiskoe) village. References to occurrence in the first half of the last century at Krasnoyarsk (Radde, 1862) are dubious. Probably the region in a broad sense is intended. 289 223 the Angara and upper Lena in the north at least to Balagansk.’ This is the northernmost point of the eastern part of the range. Toward the middle of the present century and moreover, even in the last century, the range of snow leopards shrank at some places. Thus ounces were already absent in Karatau or probably only wandered into the southern part of the range; they were absent in the Sayans (though likely intruders from the Altai into the westernmost parts), on the Tunkinsk balds (still possibly vagrant from Mongolia), and in Pri-Baikal. Ounces occurred in Pri- Baikal more as vagrants even in the eighteenth century. In the rest of the range the outlines given above are still maintained, although some peripheral portions are only sites of fairly regular intrusions.® Some false or extremely dubious views crept into literature about the distribution of the ounce, and were long maintained and sometimes still supported. The report of Pallas (1811) about the occurrence of the ounce along the Olekma, a tributary of the Lena, 200 versts from its mouth, is based on a very rare intrusion, or possibly is even a mistake (false understanding concerning tiger). Pallas obtained this information from the diary or manuscript of the elder Gmelin (‘“as I found in the rough notebooks (in adversariis) of the elder Gmelin’’). The statement of Pallas (1811) that the ounce probably occurred along the Uda River and between it and the Amur River had very large consequences.” Reference from Pallas (1811). This information is so positive that it is hardly possible to contradict the occurrence of the ounce in Pri-Baikal (Ognev, 1935). Mountain sheep evidently lived formerly in Pri-Baikal and the presence there of Siberian ibex was also possible. The range of the ounce in Siberia mostly coincides with the range of these species (see Volume I of the present monograph). ®Range according to data of Pallas, 1811; N.A. Severtsov, 1873; S.A. ee ese 1929; Selevin, 1929; Ognev, 1935; V. Skalon, 1936; Yurgenson, 1938; Levnev, 1939; Kolosov, 1939; Shul’pin, 1948; Chernyshev, 1950; Yanushevich, 1952; Sludskii, 1953; Ishunin, 1961; Stroganov, 1962; and others; and original material of V.G. Heptner. In outlining the distribution in Tuva and other regions farther east, the data of Pallas (1811) have been specifically used. According to Pallas (original in Latin), the ounce (which he called Felis pardus) **. . . intruded from Central Asia along the montane forest deserts at places in southern Siberia here and there [passim], was seen more rarely in the Altai mountains, and more often at the sources of Yenisei proper and Kemchik (Kemchug) and in the mountains around Baikal . . . On the Lena it was often caught in the vicinity of the city of Balagansk on the Гепа... A stuffed animal is preserved in the Academy Museum; the skin is that of an animal killed around Tunkinsk at Baikal’? (рр. 17 and 18). This information has been ignored or contradicted by some authors without adequate justification, mainly due to ignorance of the original text. °According to the original text of Pallas (1811, pp. 17 and 18, translated from the Latin), ‘“They say the animal was reportedly encountered (dicitur frequens esse) in the region of the Uda (Uth) River and between it and the Amur. Yakutians going there from the Lena were afraid of it and called it terrible (terribilem) to the utmost extent (ka-chai) and on encountering its tracks did not dare cross them.”’ 224 290 Some earlier workers investigating the Amur and Ussuri regions (Mid- derdorff, 1857; Maak, 1859; Schrenk, 1859; Radde, 1862; Przheval’skii, 1870; A. Nikol’skii, 1889), evidently without skins or skulls in hand, presented all their information on leopard as pertaining to ounce, relying entirely on the authority of Pallas. It should be remembered that Pallas merely passed on this information with extreme caution, prefacing it with ““They say.’’ Even after him, as quite often happens in copying from another author, the assumption turned into an affirmation (Zhitkov, 1936) and has been used as such to our time. Even some specific references to the Stanovoi and Yablonovyi ranges, which Pallas never mentioned, began to appear in the literature. From the account given above, it is quite clear that Pallas reported the mistaken accounts pertaining to tiger,!° such as the instance of an intrusion from the Olekma reported from Gmelin. Tigers in more recent periods have moved still farther north. The situation is similar in the extreme west of the range under consideration. Information about the occurrence of the ounce in the Kopet- Dag, apparently quite convincing (Khristof, 1882*; Zarudnyi, 1891*; Satunin, 1905; Bil’kevich, 1918; Ognev, 1935; and some others), and quoted extensively even in textbooks, is incorrect. A special study conducted in the Kopet-Dag (V.G. Heptner) revealed that the ounce is absent there. All affirmations are evidently based on dubious answers to queries in a study of the particularly rich and very light-colored winter skins of leopards (P. pardus). The ounce is, of course, absent on the Serakhsa (Zarudnyi, 1891*) and conditions there are not favorable for its residence. The mounted snow leopard exhibited in the Ashkhabad museum was prepared from a skin brought by S.I. Bil’kevich from ‘“Eastern Bukhara’ (V.G. Heptner). After. almost 90 years of studies in Turkmenia, not one skin or skull of snow leopard has been found. To resolve the question of the ounce in Turkmenia, it is important to note that the question of its occurrence in Asia Minor and Iran, especially in districts adjoining Turkmenia, as shown by special studies (Pocock, 1930 and 1939), have yielded negative results. The latest information for Iran (Misonne, 1959; Lay, 1967) is also so confused and indeterminate that zoologists have acknowledged the absence of proof for its occurrence in that country. In foreign literature about Iran reference is sometimes made to the data of Ognev (1935) about the occurrence of this animal in the Russian part of the Kopet-Dag as conclusive, and based on this the possibility of its existence in the Iranian Kopet-Dag assumed. It should be kept in mind !0Middendorff (1857, р. 75) similarly comments that in the case of the Tyrma, the account pertained positively to tigers. Middendorff himself found no signs of the occurrence of the ounce in the Stanovoi range. *Not in Literature Cited— Sci. Ed. 225 ро that leopards in southem Iran are particularly light-colored, almost white, especially in winter, which may have confused contemporary and past (Blanford, 1876) authors. It is significant that in Afghanistan the ounce inhabits only the extreme western, much higher parts of the Hindu Kush adjoining the Pamir (Badakhshan and Bakhan; Kullmann, 1965, 1967, and 1968). Geographic Range Outside the Soviet Union The range outside the Soviet Union (Fig. 137) covers the Mongolian and Gobi Altai, Khangai and the montane district of Lake Kosogol [Hobsogol] in the Mongolian Peoples Republic, Tibet to the Altyntagh on the north and Kam in the east, and the Himalayas from the eastern end to Kashmir. In Afghanistan this species inhabits the Hindu Kush. Here it occupies only the extreme eastern portions (see above). The leopard inhabits much lower levels there, mainly in the west and north, and relations between the two species are the same as in southern Tadzhikistan. (V.H.) 224 Fig. 137. Reconstructed species range of snow leopard, Uncia uncia’ Shreb. V.G. Heptner. Geographic Variation Ounces do not exhibit geographic variation, or else it is very insignificant. A distinct form, И. и. uncioides Horsf., 1855, has been described from Nepal, but this separation was done without adequate substantiation. Nevertheless some contemporary scientists favor recognition of the Nepalese form (Haltenorth and Trenze, 1956). On the other hand it has been shown (Pocock, 1930 and 1941) that Himalayan animals do not differ from those of the Altai. Apparently this is correct. The feature suggested as a diagnostic character (color) falls in the category of individual variation. The same applies to the form schneideri Zuk., described from a photograph of a skin (!). The absence of clear geographic variation in the ounce is due to the relatively small range of the species. It is an extremely stenotopic species and everywhere in its range is restricted to identical conditions of existence. If in spite of this a taxonomic separation of Himalayan animals is proved, the name U. и. uncia Schreb., 1775 should be assigned to the Russian form. (V.H.) Biology Population. The ounce is rare in the eastern Pamir (Tadzhikistan) but common in the western sector. They are fairly common in the Alaisk, Gissar, Zeravshan, Turkestan, and other ranges. In recent years in Tadzhikistan about 60 are trapped and caught live each year. In Uzbekistan this predator is not rare in the Ugamsk, Pskemsk, Chatkal’sk, Gissar, and Turkestan ranges, but is definitely rare in the western and southern spurs of the Tien Shan. The ounce is common in most of the Tien Shan ranges in Kirgizia. About 30 to 40 animals are captured there every year. In Kazakhstan this predator is extremely rare in the Karatau, but common in the Talassk Alatau and other ranges of the Tien Shan. Here about 10 to 15 animals are now caught annually. The animal is rare in the Tarbagarai and extremely rare in the Saur, southern (Kazakhstan) and central Altai, and also in the western Sayan and Tannu-Ola. In the latter, only solitary animals are caught and these not every year. Habitat. In summer the ounce is confined to higher levels in the majority of mountains, mostly in the subalpine and alpine zones, and often is encountered at the limit of permanent snow. In the high Alichur, in the Pamir, tracks are sometimes detected, even in winter, at a height of 4,500 to 5,000 m above sea level (R.N. Meklenburtsev). Ounces at that time are confined to mountain crests where the snow has been blown off by the wind. In the Gissar range, snow leopards live in the high mountains where rocks abound, but descend in winter into the shrub junipers along with the ibex, but not 227 293 below this level (Leviev, 1939). On the northern slopes of the Turkestan range ounces have been observed in summer only in the upper half of the juniper belt, commencing roughly at a height of 2,600 m or more above sea level. Here they are confined to rocky sites and do not leave the shrubs. Farther north, in the Chatkal’sk range, following herds of Siberian ibex in winter, the predator descends on sunny slopes from heights of 2,000 to 2,500 m above sea level to below the level of tall junipers (Korelov, 1956). In Talassk Alatau the ounce inhabits areas between 1,800 and 3,500 m above sea level. In summer, following goats [ibex] ascending to escape from blood-sucking flies and heat, it also ascends to subalpine or alpine levels; in winter, chasing goats, it descends along gorges, for example penetrating far into Aksu canyon, and visits practically all zones of these mountains. In the Aksu-Dzhebagly preserve the snow leopard is not rare in spring and early summer at lower levels along river beds and in the tree-shrub belt along slopes of gorges, since pregnant females or mothers with young goats, arkhar sheep, and roe deer which it hunts are found there at that time (Shul’ pin, 1948; F.D. Shaposhnikov, 1956; V.V. Shevchenko). In the Karatau, the ounce is confined to sites of maximum altitudes (1,500 to 2,000 m above sea level), namely, the Ker-Dzheilyau where there are many precipitous gorges in which wild apple trees, vines, dog rose and other shrubs grow, at places altogether impenetrable to man (Antipin, 1953). In the Tuyuk gorge of the Kirgiz range adjoining the Talassk Alatau, the ounce usually inhabits forestless areas of high mountains, down to the upper forest level. There they hunt goats, marmots, and snow cocks [Tetraogallus|. Hunting goats at the top of this gorge in May and June becomes difficult and is not without danger. On steep hill slopes the snow thaws intensely during the day under solar radiation; these slopes are covered either by a very stable and slippery ice crust or a thin and unstable crust. Avalanches are common at this time of year. In spring pregnant ungulates and mothers with young descend to the coniferous forest zone and dense shrubs, seeking shelter there. The leopards trail them and are regularly encountered in May and June in spruce forests where they mostly catch pregnant goats and their young, a fairly easy quarry. In the Karabaltinsk gorge in summer snow leopards sometimes descend from the high mountains into the juniper tree belt to hunt for roe deer and marmots (Spangenberg and Sudilovskaya, 1954). In the Kungei Alatau range the ounce is met with from time to time in summer in the spruce forest belt (2,100 to 2,600 m above sea level), later in the region of creeping juniper and subalpine meadows (2,600 to 2,800 m), and very frequently in the alpine level (up to 3,300 m above sea level) (В. Petrov, 1950). In the Trans-Ili Alatau and central Tien Shan snow 294 226 Fig. 138. Typical habitat of snow leopard in the Talassk range. А regular trail runs through a pass along the crest of the mountain side. Aksu-Dzhebagly preserve, western Tien Shan. May, 1955. Photograph by F.D. Shaposhnikov. leopards ascend in summer up to 4,000 m or more, sometimes descending in winter almost to “‘shelves’’ (1,200 m above sea level). In the Altai the ounce is also encountered in the alpine zone and on balds, but only at places where ibex live (F.D. Shaposhnikov). As in other regions, here too they sometimes descend along river gorges. While residing in the upper regions of high mountains, the ounce selects 228 sections with small open plateaus, gentle slopes and narrow valleys covered 227 295 with alpine vegetation, alternating with rocky gorges and jumbled rocks and debris. Siberian ibex and argali sheep live in such places in summer and sometimes in winter, and the population of snow leopard there depends on their abundance. In the central Tien Shan, it is confined to high mountain plateaus, such as elevated watersheds, to which the predator is attracted by the presence of argali sheep. Depending on the snow cover and presence Fig. 139. Kish-Kaindy gorge (2,700 m above sea level). Permanent habitat of snow leopards in Aksu-Dzhebagly preserve. Tracks of ounce (male) visible on snow in foreground. Talassk Alatau, western Tien Shan. May, 1955. Photograph by Е.О. Shaposhnikov. г» 296 of sun-warmed sites, the distribution of ungulates varies and hence that of ounce also. In general, however, in winter the ounce is confined to lower levels in mountains than in summer, penetrating the forest belt. Never- theless, as mentioned above, not only in winter but also in spring and the first half of summer this predator often lives in the tree-shrub zone because ungulates are abundant there in these seasons of the year. Yet it should be noted that the snow leopard is not a high altitude animal everywhere. There are regions where it lives year-round in low mountains and montane steppes no more than 600 to 1,500 m above sea level. As in high mountains, the predator lives here along rocky gorges, cliffs, and rock outcrops, since goats and argali sheep prefer these habitats. At a height of 600 to 1,000 m above sea level it is therefore quite common year-round in the spurs of the Dzhungarsk Alatau, i.e., in the Katu and Aktau mountains, and also in the Altynemel, Chulak, and Matai (1,000 to 1,500 m above sea level). Neither are they rare in the low spurs of the Trans-Ili Alatau and Ketmensk—the Malyi and Bol’shoi Bogut, and the Bartogoe, places covered with steppe and desert vegetation. All these hills abound in rocky outcrops among which goats live regularly, but arkhar sheep are rarer. From the foregoing description of the habitats of the ounce, one may conclude that this animal is not only an ‘‘alpine’’ animal, as commonly assumed, but also a ‘‘rock’’ species to a certain extent. Rock cliffs usually contain less snow and hiding in them when threatened is much easier; hence goats and rarely argali sheep live in such places. However, the absence of snow or a large accumulation on rocks and mountain crests gives this predator the advantage of more or less free movement during its hunting. Direct observations have revealed that the ounce is poorly adapted to moving over deep, loose snow. In areas of loose snow they regularly take advantage of beaten trails. When loose snow abounds everywhere the predator moves along sun-warmed slopes, encouraged to do so because ungulates also move there. Thin unstable-ice crust can interfere with an ounce’s hunt. Therefore, in winters of deep snow cover on mountains, ounces begin to starve and at these times approach villages and attack domestic animals more often than normal. To what extent deep snow cover impedes ounce movement can be judged from the fact that two top-notch hunters, encountering this predator on an open slope covered with deep snow, quickly overtook and captured it by throwing a chapan (top coat) over it and bundling it up. In rocks it is quite easy for an ounce to conceal itself since it has a fairly good protective coloration. Detecting an ounce hiding or creeping among rocks is very difficult. Furthermore, the snow leopard is well adapted to the severe climatic conditions prevailing in the mountains of Middle and Central Asia. Of all the big cats this species boasts the longest and most Fig. 140. Cirque of Bol’shoi Alma-Atinsk Lake on the northern slope of the Trans- - Ш Alatau, habitat of snow leopard. Eastern Tien Shan, 2,650 m above sea level. January, 1962. Photograph by A.A. Sludskii. dense hair; in addition there is hardly any difference in density of hair between winter and summer coats. In the Himalayas ounces have been sighted at heights of 5,400 to 6,000 m (Lydekker, 1924) and are usually not encountered below heights of 2,000 to 2,500 m. In summer they are seldom seen below heights of 4,000 to 4,500 т (Zherdon [= Jerdon], 1874*; Pocock, 1939). Food. The main prey of the ounce almost everywhere year-round is the Siberian ibex, and more rarely argali sheep, roe deer, young hog, and other small ungulates. In the Pamir it mostly feeds on roe deer (Rozanov, 1935; Flerov, 1935; Meklenburtsev, 1948; Egorov, 1955), attacking argali sheep occasionally. In the eastern Pamir ounces rarely attack wild sheep, especially adults. In the Gisser range, in addition to ibex, ounces hunt marmots and snow cocks (Leviev, 1939). In the central ranges of the Tien *Not in Literature Cited— Sci. Ed. 298 Shan snow leopards feed mainly оп ibex and argali sheep; quite often they catch snow cocks, marmots, and even relict ground squirrels* and mouse- like rodents (Shnitnikov, 1936; Kuznetsov, 1948), in addition to roe deer, hare, and rock partridge (Kashkarov, 1932). In the Talassk Alatau the main prey of the ounce is the ibex (Shul’ pin, 1948; Shaposhnikov, 1956). The remains of argali sheep, roe deer, piglets, and infrequently domestic sheep are found there. Sometimes in winter snow cocks and rock partridges become prey (Shul’pin, 1948). The above are usually caught at night. In the forest belt of Aksu-Dzhebagly preserve (Talassk Alatau) a two-year-old bear killed and partly eaten by an ounce was found (Fig. 144). Judging from traces of the fight, the bear was taken unawares while digging for roots in a rock stream. It tried to save itself by climbing onto a rock. The leopard had eaten the rear portion of the body and the viscera but had not touched the remainder, or the skin (F.D. Shaposhnikov, 1956). Here this predator had been lying in wait in the early morning for roe deer approaching on a solonetz. Fig. 141. Habitat of the snow leopard around Lake Issyk, Тгапз-Ш Alatau, eastern Tien Shan, August, 1960. Photograph by V.G. Heptner. *Spermophilus relictus—Sci. Ed. 299 230 Fig. 142. Low desert mountains, habitat of the ounce in Chulak mountains, spurs of the Dzhungarsk Alatau, eastern Tien Shan. September, 1949. Photograph by M.A. Kuz’mina. In Kirgizia and Trans-Ili Alatau the main prey of ounce also is ibex but quite often roe deer become victims, especially in winter (Ionov, 1929; Sludskii, 1939). At some places in the Trans-Ili Alatau roe deer are primary winter food of the ounce (B.A. Beloslyudov). The summer iair of one predator was littered with a large number of remains of ibex and two piglets. In the Dzhungarsk Alatau one often comes across goats killed by the ounce (L. Sviridov). In the low spurs of this range (Altynemel’, Chulak) the stomachs of two predators caught in November contained only remains of ibex (V.V. Afinogenov), while in other spurs (Aktau and Katutau mountains), the ounce feeds mainly on the abundant arkhar sheep, and also catches goitered gazelle (Antipin, 1941 and 1953). In the Altai and Sayans the snow leopard lives only in those ranges where ibex are common. In the Himalayas the ounce attacks ibex, gorals, wild sheep, small species of deer, Tibetan hares and marmots, and of birds—pheasants. In Kashmir they kill domestic goats and sheep, and also horses (Zherdon [= Jerdon], 1874*; Pocock, 1939). In the Mongolian Peoples Republic the main prey is ibex, more rarely argali sheep, and sometimes snow cocks and rodents. *Not in Literature Cited — Sci. Ed. | 300 In the Trans-Ili Gobi the snow leopard may descend into oasis groves to catch kulan and goitered gazelle on their way to water holes (a reported attack of this predator on adult kulan is very unsubstantiated). In the Mongolian Peoples Republic, the ounce also attacks domestic cattle but such reports are few (Bannikov, 1954). In China, in the region of Lake Kukunor [= Qinghai Lake], the remains of goats”, argali sheep, eared pheasants, and even pigeons were found around two dens of snow leopards with cubs. It is quite evident that throughout its range the ounce is almost a true stenophage since their major food nearly everywhere is Siberian ibex; argali sheep, roe deer, and other animals are eaten only in a few places. The ounce hunts ibex of all ages, but more often females and kids. Early in summer it catches mainly kids. Some summer feces of this predator were analyzed and consisted exclusively of the remains of young goats (Ognev, 1940; Shul’pin, 1948). Among the remains of 14 goat kids found in the feces of ounces in the eastern Pamir, only two contained the horny remains of Fig. 143. Remains of argali sheep (Ovis ammon karelini) killed by snow leopard ata salt lick. Aksu-Dzhebagly preserve in the Talassk Alatau, western Tien Shan. June, 1953. Photograph by F.D. Shaposhnikov. *Not clear that this is meant to refer to domestic goat, but the wild goat, or ibex, does not occur around Kukunor.—‘Sci. Ed. 233 301 hooves of kids aged three or four weeks; the remainder were those of newborn kids. Evidently ounces catch kids before they are able to follow behind their mother. From June to August they attack from time to time even adult goats 4 to 10 years of age (four instances). Many ibex are killed by the ounce in winters of heavy snowfall. For example, in the winter of 1949/1950 the remains of 25 ibex killed by this predator, including 17 males, were found scattered over a stretch of 10 km. Ounces generally kill old males weakened during rut in winter. Of 30 goats killed by ounces, 8 (27%) were females. Of the males killed, 64% were six years or older and only 36% four or five years old (Egorov, 1955). In regions where the ounce population is high ibex become scarce or disappear altogether (Koksu, Dzhungarsk Alatau—1946 and 1950; Lugovaya region, Kirgiz Alatau— 1946). The ounce rarely attacks domestic animals, chiefly sheep and goats while grazing on alpine meadows (Kashkarov, 1927; Ionov, 1929; Sludskii, 1939; and others). Sometimes the predator will sneak into a pen containing a large number of animals and kill several but a lone individual is more common. For example, in the winter of 1955 an ounce attacked a flock of sheep grazing on a winter pasture in the Koilyu valley (central Tien Shan) and killed eight sheep. In February, 1952 an ounce living in the Ozerno gorge killed seven sheep. In November, 1953 snow leopards descended very low along the Kazachka River gorge near Alma-Ata, and killed four goats (А.З. Galunshchikov). In February, 1957 in the Kosh-Agach region (Altai) one broke the roof of a sheep pen at night, but was held at bay by dogs and shot. Under exceptional circumstances a snow leopard may kill even big animals: in 1927 one killed a young horse (Ionov, 1929) and in 1938 a pair of ounces killed two colts on the northern slope of the Talassk Alatau (Kuznetsov, 1948). In winter, with the formation of a deep snow cover which makes hunting difficult, ounces sometimes approach human dwellings and attack cattle more often than in summer. In November, 1928 in the Saur an ounce invaded a sheep pen one night and killed 82 sheep. On February 11, 1937 a pair of young predators sneaked into a shed and killed a pig (Alma-Arasan, 20 km from Alma-Ata; Sludskii, 1939). In zoological gardens the daily ration for an ounce is 2.2 kg of meat (Obukhov and Shakhnazarov, 1949); in the wild it probably consumes no more than 3.0 kg of meat at one time. Home range. The dimensions of the individual territory of the ounce have not been accurately established. It often travels such long distances for a single prey that it is not possible to trace daily movement. While bypassing grazing ground of ibex or descending from the higher mountain belt into a low-lying gorge, the animal invariably sticks to the same trail, which usually runs along a ridge or along a stream or brook. Migrations 302 Fig. 144. Remains of two-year-old bear (Ursus arctos isabellinus) torn apart by two ounces. Aksu-Dzhebagly preserve in the Talassk Alatau, western Tien Shan. May, 1953. Photograph by F.D. Shaposhnikov. of the ounce were studied for several consecutive years in a section of one gorge near Alma-Ata. From fresh tracks after summer rains or in fresh winter snow it was established that the ounce traveled from the eastern slope of the mountain into the gorge, moved to another mountain, and returned some days later by the same route (lonov, 1929). In Aksu-Dzhebagly preserve (Talassk Alatau) adult animals confine themselves in winter to a particular hunting territory. The snow leopard regularly covers its hunting area, visiting the winter grazing grounds and shelters of wild ungulates known to it. The distance covered during these visits is considerable and hence a predator is generally seen a second time in a given site only after seven or eight days, 1.е., fresh tracks appear after this lapse of time (F.D. Shaposhnikov, 1956). In November to January, 1949/1950 in the Tersk Alatau, herds of ibex were confined to definite grazing sections and ounces observed there. At one place after a snowfall a leopard was regularly seen around an active apiary (Zimina, 1953). Its visits were evidently due to scarcity of food. 234 Fig. 145. Snow leopard chasing ibex (Capra sibirica). Broken line in upper right part of photograph denotes ounce track, and in left bottom part tracks of a group of mountain goats which had crossed that place 10 minutes before. Dark spots on slope are bushes of dwarf juniper Juniperus). Talassk Alatau in Aksu-Dzhebagly preserve, western Tien Shan. February, 1961. Photograph by E.N. Matyushkin. The present population density of snow leopard can be estimated from the fact that around the 1950’s about six to eight animals were counted in an area of about 40,000 hectares in the Aksu and Dzhebagly River valleys of Aksu-Dzhebagly preserve (Talassk Alatau) (F.D. Shaposhnikov, 1956). Burrows and shelters. The ounce usually sets up a den in a cave or crevice among rock piles under an overhanging cliff or some similar place where concealment during the day is guaranteed. Occasionally they rest in daylight hours in a sparse spruce forest, among juniper shrubs, in talus slopes, or individual rocks. In the Kirgiz Alatau the ounce has reportedly utilized the large nests of black vultures [Aegypius monachus] situated among low junipers for a diurnal bed. The amount of the predator’s fur found in such nests indicates that it must spend a significant part of the day in them (Spangenberg and Sudilovskaya, 1954). The ounce often uses the same lair for several consecutive years. Dens with cubs are always littered with bones of ibex and other ungulates. 233 304 Fig. 146. Track of snow leopard (male) in packed snow. Aksu-Dzhebagly preserve in Talassk Alatau, 3,000 m above sea level. Western Tien Shan. May, 1955. Photograph by F.D. Shaposhnikov. Daily activity and behavior. The ounce becomes active at twilight, hunting before sunset and again at dawn. It rarely hunts during the day. An ounce was observed chasing argali sheep at 9.00 a.m., and twice roe deer were found that had been killed by these predators at midday (N.A. Yanushko). The ounce has often been encountered during the day but was usually sleeping or resting on a rock. In the southern part of its range, in the Himalayas, the predator emerges to hunt even before sunset (Pocock, 1939). Captive animals lie in a corner of their cage throughout the day and begin to move only after the onset of twilight (Fig. 145). The ounce hunts by two methods: either the predator creeps toward its 235 305 prey from behind cover or lies in wait near trails leading to a water hole or salt lick, hiding on a cliff, in a precipice, or between rocks. When the prey is just a few tens of meters away, the snow leopard overtakes it in huge bounds of 6.0 or 7.0 т and attacks it quickly."' If it does not catch the prey in the first attempt, it usually desists after a few leaps. Such a method of hunting is illustrated by the following observation. ‘п August, descending into the valley from Tom’ pass, I stopped for the night. The sun had already set and it was twilight in the gorge. From all sides the calls of snow cocks could be heard. Looking at the opposite side of the gorge, I saw a herd of ibex grazing quictly a few hundred meters from where I was standing, and two ounces, a large one and a small one, evidently a mother and cub, the animals, like shadows, crept between the rocks, using the smallest as cover. On coming close to the herd, the adult ounce with an immense leap of 6.0 m jumped at an animal but missed. After two more bounds after the escaping ibex it returned and went away along the gorge accompanied by the cub’’ (Sludskii, 1939). Reports that the bounds of ounce sometimes measure up to 15 m (lonov, 1929) are poorly supported. Having caught an ibex or roe deer, an ounce throws its prey to the ground and, pinning it with the forepaws, rips open the abdomen or throat with its teeth. ‘Ап eight-year-old goat which died almost before my eyes had deep wounds on both sides of its neck, inflicted by the claws of the predator. Death was due, however, to the deep wounds in the abdomen through which the ounce pulled out the viscera with its teeth’? (Egorov, 1955). In addition to concealment, the ounce catches prey from an ambush, waylaying them from a cliff or among rocks. In the early spring of 1955 in Aksu gorge of the Talassk Alatau, an ounce was seen lying in ambush on a large stone overlooking the trail along which female ibex went to a water hole (Shaposhnikov, 1956). Judging from tracks in the Dzhungarsk Alatau the ounce often attacks ibex from an ambush, leaping from above (A. Sviridov). Sometimes this predator attempts to chase its quarry. Thus, in the Dzhebagly range ounce tracks behind a female arkhar sheep were found for a distance of one kilometer (Shaposhnikov, 1956). In mid-June, 1952, at a height of about 4,000 m above sea level, an ounce was seen frightening argali sheep and chasing them for some time (N.A. Yanushko). Unlike the wolf, during a hunt the ounce does not attempt to kill several animals at one time and is usually satisfied with one. At the end of summer, autumn, and in early winter, the snow leopard often hunts in families of two to six animals, more often two or three. For ‘1 According to Pocock (1939), as an adaptation to hunting by concealment in rocks, the eyes of ounce are set so high that while creeping up to a quarry the predator can see it without lifting its head. 306 example, in the early winter of 1954 in the Aksu River canyon (Talassk Alatau), five ounces belonging to a single family were found attacking a young hog. The predators fell on the prey almost simultaneously and quickly tore it to bits (F.D. Shaposhnikov, 1956). Having killed its prey, the leopard generally shifts it beneath a juniper tree or rock before feeding. The remains of a prey are usually left behind, and are consumed by vultures, bears, or wolves. Only on occasion does the predator stay near its kill and chase vultures away. The ounce is not a cautious animal and on noticing or encountering man does not hurry to cover. In the early morning of April 24, 1954 in Kish- Kaindy gorge (Aksu-Dzhebagly preserve in Talassk Alatau), a preserve Fig. 147. Tracks of a female snow leopard on slushy snow. Pass toward Aksu River (3,200 m above sea level) in Aksu-Dzhebagly preserve. Talassk Alatau, western Tien Shan. May, 1955. Photograph by F.D. Shaposhnikov. 236 307 employee heard his dogs barking. Rising from his seat he saw ап adult ounce just a few meters away. The animal stood in a tense pose with its tail raised and the forepaws pressed firmly to the ground. On noticing the man, it turned toward him and snarled loudly. The animal was shot and proved to be a female (F. Shaposhnikov, 1956). At the same time, the ounce is extremely timid toward humans and even when wounded will attack the hunter only in exceptional circumstances. Animals which fall into traps are usually taken alive by hunters (see below). The timidity of this predator may be judged by the following examples. Late in Summer an ounce entered a sheep pen high in the mountains on a dzhailyau*. On hearing the commotion in the pen, the shepherdess rushed in and, catching the animal by its tail, with shouts began to drag it away from the wounded sheep. The ounce did not attempt to protect itself and was killed by shepherds who had arrived on the scene. In the Kara-Kurdzhur region of the Tien Shan dogs once attacked an ounce approaching cattle. Taking advantage of the attack, a young shepherd threw his chapan [coat] over it and bound the predator (Voinov, 1956). Instances are known of a captured adult ounce which allowed itself to be stroked after several days and even permitted itself to be approached inside its cages. Taken young, ounces become fully tame in a very short while. Fig. 148. Trail of an ounce in loose snow in Aksu Dzhebagly preserve. Talassk Alatau, western Tien Shan. February, 1961. Photograph by G. Kuznetsov. *Kazakh word meaning “‘high pasture’’— Sci. Ed. ta > a Only two instances are known of ounce attacks. on humans. On July 12, 1940, in Maloalmaatinsk gorge near Alma-Ata, an ounce attacked two men during the day and inflicted serious injuries on both. Investigations after the animal was shot established that it was rabid. In the second case, not far from Alma-Ata, an old, toothless, highly emaciated animal jumped from a rock onto a passerby in winter. It was stunned by a blow from a stick, tied up, and carried to the village (A.A. Sludskii). Seasonal migrations and transgressions. These are poorly known. Regular vertical movements while following the trail of ungulates are clearly known. In summer part of the population moves into the subalpine and alpine belts and in winter, with the onset of deep snow, moves into the forest zone and even farther below (see ‘‘Habitat’’). An instance is known of an ounce from the southern part of Lake Telets ascending into the upper Kyga and reaching Erinat in the Bol’shoi Abakan basin (Kolosov, 1939). Such migrations are evidently associated with a search for sites rich in game. In looking for food, the ounce enters the Karatau and the spurs of the Trans-Ili and Dzhungarsk Alatau. From time to time in search of prey or to escape from deep, loose snow cover, the snow leopard descends from the mountains and occurs not only in foothills, but also in adjoining plains. In February, 1954, a leopard was killed in the foothill steppes (about 700 m above sea level) at Sartoga in the Uigursk region of Alma-Ata district (Anurov, 1954). In January, 1958, another was caught on the plains near the mountains in the Sairamsk region of Chimkent district; the animal was lying among reeds by the side of a small lake. In March of the same year three ounces (adult female and two cubs) were killed among reeds in the floodplain of the Issychka River (about 600 m above sea level), roughly 60 km east of Alma-Ata. The animals had followed hogs migrating from the mountains because the latter had very deep snow cover. In July of the same year an ounce was caught at Dzhaison in the Chuisk region of Dzhembul district, 80 km from the nearest mountains. It lived in a hilly area and was killed while attacking sheep (Tleukulov, 1958). In the winter of 1957/1958 an adult ounce was found very far away from its regular habitat and caught in a hilly area on the northern shore of [Lake] Balkhash, 60 km east of the city of Balkhash. The predator could have penetrated there from the west through the Chu-Ilisk mountains or from the east from the Dzhungarsk Alatau. In either case it traveled more than 600 km from the nearest mountains (I.G. Shubin). The appearance of ounce in very low montane zones, in desert plains, and in oases groves, where they are confined to rocks, has been recorded in the uninhabited regions of the Trans-Altai Gobi in the Mongolian Peoples Republic (Murzev, 1948). In this same country, in the Nemegetu depression of the southern Gobi, the 309 Fig. 149. Single track of a snow leopard and its tracks in different gaits (walking and bounding). Aksu-Dzhebagly preserve, Talassk Alatau, western Tien Shan. February, 1961. Sketch by E.N. Matyushkin. ounce lives on low argillaceous and sandy cliffs rising to heights of only 100 m (Efremov, 1956). Reproduction. This is not well studied. According to observations in zoological gardens, where this predator sometimes breeds, sexual maturity in the female sets in at the age of two to three years. In captive animals estrus occurs from March to May. In Uzbekistan estrus in leopards occurs from the end of February to March (P.F. Ryplya). During the breeding periods with the onset of twilight characteristic loud cries similar to bass meowing are emitted by the ounce, generally while standing on projecting rocks. The duration of gestation is 93 to 110 days (Dzhennison, 1929*; Shereshevskii, 1940). In the central Tien Shan young are born around mid-May. A den with newborn kittens was found on May 15, 1913. Sometimes litters are seen even later. Five litters in June and one in July have been recorded (Shnitnikov, 1936). In the Narynkol region, in the central Tien Shan, however, a female pregnant with two fetuses was caught at the end of March, 1957 (2.1. Bibikov). In Kirgizia whelping occurs in May and June (Kuznetsov, 1948; *Not in Literature Cited — Sci. Ed. to г 310 Berens, 1957). Early in June, 1955, in the central Tien Shan a litter of five blind kittens was found. A den was discovered among rocks only 500 m away from a camp of geologists (V.M. Katkov). In June, 1949, a female was caught with nurslings in the spurs of the Trans-Ili Alatau. In the Talassk Alatau, in Chepta gorge, ounce kittens the size of domestic cats were found at the end of May (Shul’pin, 1948). In these same mountains, in a forest of juniper trees at a height of 1,800 m above sea level, on April 24, 1954 a pregnant female with one well-developed fetus was caught; the body length of the fetus (female) was 22 cm, tail 12 cm, and weight 393 g. The fetus already had well-developed pigmented pelage. The mammary glands of the captured female were enlarged (F.D. Shaposhnikov, 1956; V.V. Shevchenko). In this case parturition would have occurred at the end of April or in the first few days of May. A litter of two blind kittens was collected in the Garm region in Tadzhikistan on May 7, 1940. In the Mongolian Peoples Republic ounce kittens —two, three, and rarely one—are born in May and early June. By the end of July kittens are one-third the size of adult animals (Bannikov, 1954). Judging from the dates of encounter given above, kittens are usually seen in May in the northern half of the range; more rarely parturition takes place in the second half of April or in June. Reports referring to the appearance of kittens in the Narynkol region (central Tien Shan) in March and early April, and in Kirgizia in early April, require confirmation (Sludskii, 1939; D.P. Dement’ev et al., 1956). A litter may consist of one to five cubs. Even larger litters are possible in exceptional circumstances; litters of seven cubs have been recorded (October, 1954; Dzhety-Oguzsk region, Issyk-Kul district; Berens, 1957). Two or three cubs are more common. In the central Tien Shan the ounce usually produces two cubs, generally a male and a female. A litter of three kittens with a single male has been observed only once (Shnitnikov, 1936). In Kirgizia an ounce may produce two to five cubs, more often two or three (Kuznetsov, 1948; D.P. Dement’ev et al., 1956; Berens, 1957). In the Talassk Alatau this cat gives birth to two or three kittens per litter (Shul’pin, 1948). Of seven litters discovered in the Trans-I]i Alatau, one contained a lone cub, four had two, one had three, and the seventh had five (A.A. Sludskii). In the southern Altai families of ounce consisting of two or three animals are observed repeatedly (Selevin, 1929). In the Kukunor district of China two dens were found, each housing one ounce and a single cub, and in Tibet a litter consisting of three cubs was discovered (Pocock, 1939). Among ounces breeding in zoological gardens two to four cubs are common (Shereshevskii, 1940). Growth, development and molt. Cubs are born helpless, blind, and with their ear passages closed. Measurements of blind cubs (May 7, vicinity of 238 Sulit Fig. 150. Fetus of a snow leopard a few hours before birth. Aksu-Dzhebagly preserve. Talassk Alatau, western Tien Shan, April 24, 1955. Photograph by F.D. Shaposhnikov. Garm) are as follows: male—body length 29.0 cm, tail 15.5 cm, and weight 430 g; female 28.0, 15.5 cm, and 460 g respectively. One male born in a zoological garden had a body length of 237 mm, tail length of 168 mm, length of hind foot 54 mm, height of ears 18 mm, and weight 440 g; the corresponding measurements for a female were: 230 mm, 181 mm, 53 mm, 17 mm, and 495 g (Bree, 1965). The pelage is very dense but quite bristly. The main shade of the coat on the back in light brown, especially in the region of the shoulders and croup. The coat throughout the upper part of the body is mottled with dark brown spots about 1.0 cm in diameter. On the neck and withers these spots are even smaller. On the croup the spots fuse into two broad longitudinal stripes between which some five or six spots are scattered. Around the lip, on the cheeks, and around the whiskers, dark, 312 almost black markings occur. The entire underside of the body, commencing from the lower lip and up to the genitalia is creamy-white. Around the upper part of the neck а single row of dark spots forms a neck band. Rarely, spots the size of peas are seen on the upper part of the abdomen. The outer surface of the tail has dark brown spots. Both the fore- and hind limbs are light in color. Even in early July the young begin to trail behind their mother. A female and two young ounces were encountered in Mynzhilka region (3,000 m above sea level), 20 km from Alma-Ata on July 10, 1937. In July, 1949 another female with two cubs was met with at Aksai. During July and later cubs play together in the mornings just like domestic kittens. Even while suckling, cubs take part in the hunt, following their mother. In November, 1948, at Bartoga (Trans-Ili Alatau) a family consisting of a female with five young was observed hunting ibex. The young were half the size of the adult. At Lake Iskanderkul’ in Uzbekistan at the end of February, 1953, a female was caught and the two cubs with her were only slightly smaller than she (Ishunin, 1961). The young remain with the mother until she comes into estrus again. The male plays no role in the raising of cubs. Longevity of the ounce is not known. Some have survived for 15 years (Pidoplichko, 1956). Molt has not been studied. A gestating female caught in the Talassk Alatau on April 24, 1954, had not yet undergone molt (F.D. Shaposhnikov, 1956). Ounces molt twice a year. The summer coat differs very little from the winter coat in density and length, and this difference is less than in other cats. Enemies, diseases, parasites, mortality, and competitors. Apart from wolf, the ounce evidently has no enemies. Wolf and leopard are serious Fig. 151. Snow leopard cub, 1.5 months old. Vil’nyus Zoological Garden. Photograph by B.B. Marm. 240 313 competitors in the southern parts of the range. Diseases suffered by the snow leopard have not been studied thoroughly. Instances of rabies are known among wild ounces (see above). In zoological gardens young animals quite often die from infectious enteritis, a highly contagious disease with a very severe course; mortality may be as high as 70 to 80% (Tsvetaeva,1949). Mites, Haemaphysalis warburtoni, were found in an ounce caught in the vicinity of Alma-Ata on October 29, 1955 (G.B. Ushakova). Population dynamics. In some years the ounce population is observed to increase sharply in one or another region. Comparatively large numbers were sighted in 1946 in the region of Lugovaya station, Kirgiz Alatau, and in the Gvardeisk region, Dzhungarsk Alatau. In the latter region ounces were numerous in the winter of 1949/1950. In the winter of 1952/1953 an increase in snow leopard population occurred in Kaskelensk region of the Trans-Ili Alatau mountains and in the Taldy-Kurgansk region of the Dzhungarsk Alatau; in the latter region a simultaneous rise in the lynx population was also noted. In the following winter of 1953/1954 the ounce and lynx population increased considerably again in the Dzhungarsk Alatau and even in the Panfilovsk region (V.S. Korovets; M.S. Kuznetsov; and others). It is possible that this increment in population of both predators was due to an abundance of ibex in these areas. In general, factors controlling changes in the population of the ounce are not known. In the 1940’s to the 1950’s a reduction in hunting led to an increase in snow leopard stocks everywhere. Field characteristics. The ounce is a large cat of smoky-brownish-gray color mottled with large, annular, dark brown spots. The tail length exceeds three-fourths of the body length; it is densely furred and appears even longer and thicker. Tracks are large, rounded, and without claw impressions. Remains of ibex killed by the ounce are usually seen close to steep cliffs and rock piles, under juniper, and in other such sites; wolves kill them only on gentle slopes, far away from rock outcrops. The remains of prey of the ounce and wolf differ considerably in the following respects. In the skull of an ibex or arkhar sheep killed by ounce, the nasal bones and orbits remain undamaged. Around the skeleton of an ibex killed by the ounce invariably one finds large bits of skin. The wolf, contrarily, generally punctures the nasal region of the skull, crushes the orbits, punctures the lower part, and frequently enlarges the foramen magnum in four- to five-year-old arkhar sheep and ibex to extract the brain. The wolf strips the skin into small pieces or eat it completely. Even if a wolf has scavenged the remains of the leopard’s prey, the fact of the latter’s kill can still be ascertained from the large scraps of skin scattered around. When the death of an ibex or arkhar sheep is not due to these two predators, vultures quickly find the carcass and having eaten the carrion, turn the skin 240 314 Fig. 152. Snow leopard оп cliff in the Тгапз-Ш Alatau, eastern Tien Shan. Photograph by M. Galkin. inside out; they do not tear it to pieces as done by the ounce (Egorov, 1955). (A.S.) Practical Significance The ounce is of little importance to the fur industry. In the 1950’s to the 1960’s only a few tens of skins were tanned everywhere. Consequently their tanned value was very low, on the average about 3 rubles. The main regions of ounce hunting are Tadzhikistan and Kirgizia. However not long ago in Kirgizia, up to 60 skins were prepared annually. Thus, in 1925/1926, 37 BIS skins were prepared, 1926/1927—56 skins; 1927/1928—20 skins; 1928/1929—37, 1931—53, 1936—54, 1937—45, 1938—31, 1939—45, 1940 —27, 1942 —16, 1943 —24, and 1944— 13. In the 1950’s and 1960's, 6 to 18 skins were tanned annually in Kirgizia. In Kazakhstan in recent years fewer than 10 ounce skins have been tanned. A larger number were tanned there formerly. In 1926/1927, 8 skins were prepared, 1928/1929—66, 1933—4, 1934—15, 1935—20, 1936—3, 1937—3, 1938—13, and 1939—14. More skins of snow leopard were prepared in Tadzhikistan in the 1950’s: 1951 —16, 1952—8, 1953—17, 1954—27, 1956—35, 1957—33, and 1958—S3. In Uzbekistan over the last 10 years, 3 to 8 skins were tanned annually, mainly from Surkhan-Dar’insk regions: 1951—5, 1952—6, 1953—6, 1955 —6, 1956—3, and 1957—3. It is thought that presently about two- thirds of the skins of snow leopard remain with the people. Some lone ounces have been caught in Altai and the Sayans. For the Soviet Union as a whole, 118 ‘Чеорага [bars]’’ skins were tanned in 1928/1929; of these, 8 were prepared by the Siberian group of firms, and 110 by the Middle Asian group (Kogan, 1931). This number also included skins of common ‘Чеорага’” [/eopard] and cheetah, comprising about 20 skins. In the Soviet Union in the 1920’s some 100 skins of ounce were tanned annually: 1926/1927—70 skins, 1927/1928—30, 1928/1929—120, 1931—82, 1953—29, 1954—52, and 1956—120 (Kuznetsov, 1932). In the last century ounce hunting was intensified and in the Semirech’e district up to 140 caught annually. The magnitude of hunting can be judged from the following data on dressed skins (Silant’ev, 1898; Shostak, 1927): 1884—83, 1885—29; 1886—142, 1890—37, 1891—54, 1892—106, 1893 —47, 1894—74, 1895—36, 1897—58, 1898—76, 1899— 29, 1900—30, 1901—36, 1903 —47, and 19905 —70. Of these, the bulk came from the Przheval’sk and Pishipeksk sections of Киса. Kazakhstan provided 10 to 20 skins (Dzharkent section). The annual output of ounce skins from the Soviet Union, compared with the world output, was very much greater in the past. From 1907 to 1910 the annual world output of snow leopard skins was placed at 750 to 800. In some years up to 500 leopard skins passed through Nizhnegorod market, while the world output was estimated at 1,000 (Zhitkov, 1928; Kuznetsov, 1932). Thus Russia formerly contributed up to 50% of the world output of ounce skins. Mongolia produced 80 skins in 1908, 40 in 1927, 40 again in 1934, and 15 to 25 annually until 1944 (Bannikov, 1954). All the 120 leopard skins put up for sale in 1956 at the Leningrad auction were sold to the USA, at an average price of $11.25 per skin. These skins were mainly used in making carpets and fur coats for women. 316 Early in the present century methods of catching live ounces were developed in Kirgizia, and catches either exported or placed in Soviet zoological gardens. Catching ounces alive is practiced at present in Kazakhstan, Kirgizia, and Tadzhikistan. Live animals are handled by Zootsenter. From one, the Issyk-Kul’ district (Kirgizia) alone; this organization in the 1950’s received 50 ounces in three years (Berens, 1957), while 14 live animals were captured in the first half of 1958 in all of Kirgizia. During the 1940’s and 1950’s the Przheval’sk branch of Zootsentr in Kirgizia received 375 live leopards, or an average of 18 animals per year. The Tadzhikistan branch of Zootsentr, located in the eastern Pamir, received 50 leopards in 10 years (1950’s). In individual years the organization handled up to 10 animals. In the world market the ounce is always in great demand and expensive. Within the Soviet Union a live animal is priced at several hundred rubles. Though the snow leopard became known in Europe even at the end of the eighteenth century, they first became acquainted with a live animal only in 1872 when Governor-General Kaufmann sent a pair of young animals from Turkestan to the Moscow Zoological Garden. The damage caused by these predators to animal husbandry in Kazakhstan is of no practical significance. In regions where livestock 242 Fig. 153. Snow leopard taking refuge in a tree. Vakhana gorge, Pamir. Photograph by G. Lipskerov. ST are many and the ounce common, no more than one or two attacks have been reported. Mainly stray sheep and goats become victims (Sludskii, 1939). They cause little damage to stock even in Kirgizia where the predator is particularly common (Kuznetsov, 1948; D.P. Dement’ev et al., 1956). They rarely attack man, even when wounded. Only two attacks are known— one by a rabid ounce and another by an extremely emaciated old animal (see above). Live ounces are caught by placing large traps woven from sheep’s wool or loops of steel cable along their trails. Loops are usually placed together with a small trap with weak springs. Having set up the trap, the loop is placed on top of it and the free end tied to a spring on a strong and resilient pole. An ounce having stepped into a trap, simultaneously is grabbed by the loop. The latter is quickly retracted by the spring, hoisting the animal into the air by its paws in such a way that its hind feet can still touch the ground. To ensure that the loop does not injure the animal’s foot, a restrictor is first attached which prevents excessive tightening. An ounce on falling into a trap usually executes one or two leaps, then sits quietly. It cannot escape even if only one or two digits are caught in the trap. Sai Fig. 154. With a captured snow leopard. Tien Shan. March, 1959. Photograph by A. Grecher. 318 The ounce is removed alive from the trap or loop in the following manner: one hunter stands in front of the animal and distracts it while another approaches from the rear and throws an overcoat or chapan over it in order to catch it by the head. The paws are then bound together and tied down, a stick is placed between the teeth (tied together), after which the animal is released from the trap or loop. Sometimes a trapped animal is noosed at the neck, one paw slipped into a loop tied to a beam (‘‘ikryuk’’), and then the animal trussed. Often hunters approaching a captured animal adopt a different procedure. One man moves forward to distract the ounce or places a stick in its mouth, while another at an appropriate moment catches its tail and drags the body toward himself with force. The head of the dragged animal is quickly covered with an overcoat, padded jacket, or a thin felt material made of sheep’s wool so as to temporarily blind it. A blinded animal quiets down and can be tied up easily. Bound and gagged, the animal is then strapped to the saddle of a horse, carried on the back (young animal), or pulled on a sledge of branches to the camp and placed in a temporary cage. At present, several hunters are known who have successfully trapped ounces by the methods described above, among them a widely known hunter living in the Issyk-Kul’ district (Kirgizia) who has trapped up to 30 of these cats (Berens, 1957), and another famous leopard catcher who caught 60 in Tadzhikistan (in Murgab region). From time to time ounces fall into traps intended for ibex or argali sheep. On these occasions they are shot. After the first snowfall ounces are sometimes hunted with the help of dogs; moreover, at the beginning of this century they were sometimes poisoned with strychnine. | In the standard fur terminology used throughout the Soviet Union ounce pelts are called ‘Чеорага” [bars]. They go into carpets, sledge covers, women’s overcoats, and more rarely fur coats, collars, etc. For a long time, due to misunderstanding, the ounce was considered a dangerous and harmful predator and hunting thus permitted year-round by any method. Rewards were even offered for dead animals. In reality, as stated before, the harm done to livestock and game is insignificant, and the cat poses no danger whatsoever to man. The ounce is, in fact, a valuable treasure in our mountains and of great scientific interest as a highly specialized wild cat. In the world market live ounces are always in demand, and their sale a profitable item for zoological exports. Evidently the indiscriminate destruction of this animal should be banned everywhere. Catching ounces should be prohibited year-round in the Tarbagatai, Saur, Altai, and Sayans. In Tadzhikistan, Uzbekistan, Kirgizia, and Kazakhstan shooting ounces should be banned and catching them for the purpose of export only permitted from October through February. In 1950 in Kirgizia, the ounce was struck from the list of predators to 244 245 319 be exterminated. Hunting was banned altogether and only the catch of live animals permitted. In this same republic capturing cubs and destroying dens with cubs are prohibited. (A.S.) Genus of Small Cats Genus Felis Linnaeus, 1758 1758.! Felis Linnaeus. Syst. Nat., 10th ed., vol. 1, p. 41. Domestic cat, D2: 1821. 1829. 1834. 1841. 1843. 1855. 1858. 1858. 1858. 1858. 1867. 1898. 1903. 1905. 1926. 1.е., Felis libyca Forst. Lynx. Kerr. Anim. Kingd. Cat. Mamm., NN 288-289. Felis lynx Linnaeus. Lyncus. Gray. London Med. Rept., vol. 15, p. 302. Felis lynx Linnaeus. Pardina. Kaup. Entw. Gesch. Nat. Syst. Europ. Thierwelt, vol. 1, pp. 53 and 57. Felis pardina Temm. = Felis lynx Linnaeus. Lynchus. Jardine. Nat. Libr. Mamm., vol. 4, p. 274. Felix lynx Linnaeus. Otocolobus. Brandt. Bull. Sc. Acad., St. Petersb., 1, 9, 38. Felis manu! Pallas. Caracal. Gray. List. Mamm. Brit. Mus., p. 46. Caracal melanotis Gray = Felis caracal Schreber. Catus. Fitzinger. Wiss. Popul. Nat. d. Sdugeth.,vol. 1, p. 205. Felis catus Linnaeus (domestic cat). Catolyx. Severtsov. Revue. Mag. Zool., vol. 10, p. 387. Felis chaus Giildenstaedt. Prionailurus. Severtsov. Ibid., р. 387. Felis pardochrous Hodgson = Felis bengalensis Kerr. Urolynchus. Severtsov. Ibid., р. 389. Felis caracal Schreber. Oncoides. Severtsov. Ibid., р. 390. Given to one of the South American cats but applied to euptilura Elliot (Satunin) with a reference to Severtsov. Cervaria. Gray. Proc. Zool. Soc. London, p. 276. Lyncus pardinus = Felis lynx Linnaeus. Oncoides. Trouessart. Cat. Mamm., vol. 1, p. 357. Nomen praeoccupatum (Severtsov, 1858). Eucervaria. Palmer. Science, N. S., vol. 17, p. 873. Replacement for Cervaria Gray. Trichaelurus. Satunin. Ezheg. Zool. Muzeya I. Ak. Nauk, vol. 9, p. 495, St. Petersb. Replacement for Otocolobus Brandt. Eremaelurus. Ognev. Ezheg. Zool. Muzeya I. Ak. Nauk, vol. 27, ‘Some of the names given were coined for subgenera, but also used as generic names. oe) i) > р. 356, St. Petersb. Eremaelurus thinobius Ognev = Felis margarita Loche. (V.H.) Small- and moderate-sized cats; a few species are large. The general outward appearance is typical of small cats, 1.е., sacral region high and stands above shoulder, and back arched forward; more rarely, back line straight (see Fig. 30). Legs mostly short or moderate in length, as a result of which the animal appears long; a few are long-legged and these animals appear nearly squarish. Tail varies in length; in a few species very short, not more than one-seventh of body length (trunk with head), and does not reach calcaneal joint in standing animal; in most species of moderate length, varying from one-third to one-half of body length; and in a few species equal or almost equal to body length. Head relatively large, short and broad in most, almost spherical in some, slightly elongated in a few, and perceptibly so in some, resembling somewhat the head of big cats (genus Panthera; puma and clouded leopard). Ears vary in size and shape but almost invariably relatively larger than those of big cats (Panthera) and ounce (Uncia), and prominently project beyond even the winter coat. Usually they are quite large and pointed but sometimes blunt; sometimes they are very large and steeply set; or they may be pushed sideways (‘‘overhung’’) and the parietal region between them very broad. In some species the hair on the top of the ear is only slightly elongated; sometimes, however, the long pointed ear has a well-developed tuft of long hair at the tip. Eyes relatively large and very large in most species; pupil usually vertical but round in some. Hair coat uniform throughout the body; hair on cheeks in a few species long and forms fairly well-developed whiskers. The feet in most species are not strong and the paws relatively small; in a few big and moderate-sized species, they are strong with powerful, broad paws similar to those of ounce and big cats (puma, lynx, and clouded leopard). The claws in all species except one (F. planiceps, Kalimantan) are wholly retractile and their ends do not project out of the fur. Skin folds and interdigital membrane quite variable, indicating various degrees of development of this adaptation. In some species (clouded leopard, marbled cat—F. marmorata, and others), interdigital membrane and skin folds over claws fully developed, i.e., folds are doubled up, occur on all the digits, and the membrane reaches ends of digital cushions (as in most cats and ounce; see Fig. 278). In many species the skin fold is the same size on each digit in both fore- and hind paws, and covers retracted claws of digits Il and Ш from the inside and of digits ТУ and У from the outside, while the interdigital membrane is relatively poorly developed. In some species skin folds cover the claws but the interdigital membrane is almost undeveloped (caracal, similar to cheetah; see Fig. 278).? Description of paw structure here and elsewhere taken from Pocock (1941). 246 ЭТ Pelage usually dense, soft, silky, and relatively long, especially in winter. Color highly variable, from wholly monochromatic, often yellow shades, to monochromatic with traces of spots, usually on the extremities, of various color shades. Spots highly variable in size, ranging from blotchy to annular to small, or small fused into large ones. Often these spots form bands or fuse, or almost so, into longitudinal and more rarely transverse small stripes. A true ‘‘tiger’’ color does not occur. In some species (clouded leopard, group Prionailurus) a white spot occurs on the back of the ear as in some big cats. A few species are polyphasic (Е. temmincki and Е. yaguarundt); melanistic and erythristic individuals occur in many species and are sometimes geographically localized (leopard cat and manul). Teats, two to four pairs. Skull of most species relatively large, light in build, and with relatively thin bones. Zygomatic arches weak, orbits large and closed or almost closed in some species, and cranium large and bulging. Upper line of profile highly convex, fairly steep, and arcuate; interorbital region raised and profile, both behind and in front of interorbital region, dips steeply. Skull broad posteriorly; zygomatic width may exceed one-half of condylobasal width. General skull features viewed dorsally appear elliptical to varying degrees with a slightly elongated cranial part. Exceptionally, skull appears rounded. Crests and tuberosities absent or only weakly developed in most species; sagittal crest totally absent or seen only posteriorly in the form of an insignificant elevation; lambdoidal crests better developed but still weak. Postorbital constriction weakly defined. Facial part of skull relatively small and short, usually shorter than cranial part (reckoned from middle of supraorbital processes). Small pit (depression) present between nasal and frontal regions. In most species the gap between the posterior wall of the glenoid fossa (postglenoid process) and the tympanic bulla is very small. In a few the bullae are shifted significantly behind, or contrarily, set so far forward that the anterior margin of the tympanic bulla falls at the level of the glenoid fossa. Tympanic bullae per se very large, relatively much larger than in big cats and ounce, and bulging. Usually the entotympanic chamber is larger than the ectotympanic; in some species, however, the latter is equal to or only slightly smaller than the entotympanic chamber. The third upper element (epihyale) of the hyoid apparatus (see Fig. 31) is completely ossified, and depressions in front of the jugular foramen are absent on the basioccipital part of the occipital bone. Only in a few species is the skull relatively long and coarser, with strong zygomatic arches, well-developed crests, and a more powerful postorbital constriction, giving it the general appearance of the skull of big cats (genus Panthera; clouded leopard, F. nebulosa, the skull of which resembles that of puma, 322 Е. concolor; and to а lesser extent several other species, for example Е. temminckt). Dentition in most species is complete although the second upper premolar (first in the series) is very small. Some individuals in certain species lack this tooth; in other species, often distant from one another, the general absence of this tooth is designated a specific characteristic in spite of the fact that it is present in some individual animals. The dental formula in this genus may there be given as: 3 ее р шо 3 1 2 1 2 or [a ees pm gm eee 3 1 2 1 As an abnormality a rudimentary second lower molar is sometimes seen (lynx; Zoological Museum, Moscow University). Relatively weak teeth have also been noted in which the talon of the upper carnassial tooth was greatly reduced in size and the canines relatively thinner at the base, more slender, and relatively longer than in big cats and ounce. Particularly long canines (longest in the family) have been recorded for clouded leopard. Sexual dimorphism is absent but females average smaller. Age-related dimorphism is manifested in several species, mainly monochromatic or weakly spotted ones. Kittens have either well-developed (puma) or only faint spots throughout the body or only in some parts (extremities). Seasonal color dimorphism is absent or the winter coat perceptibly lighter than the summer one (lynx). In most species the winter coat is denser and more luxuriant (longer) than the summer coat. These are predators that prey mainly small vertebrates, mostly rodents and birds. In other words, the quarry is usually much smaller in size than the predator. Some large-sized species (puma and lynx) catch hares and animals even bigger than themselves (deer). They hunt by ambush. Members of the genus inhabit all types of terrain except tundra and occupy highly diverse biotopes ranging from high latitudes in mountains to sandy deserts. Due to the specialized structure of the hyoid apparatus (see Fig. 31) and the poor mobility of the larynx, these animals cannot produce a low ‘‘тоаг- growl’. Instead their call is very high pitched shriek, resembling that of domestic cats, and varies widely in different species (‘‘meowing cats’’). Well-being and contentment are expressed by purring during both inhalation and exhalation. By and large cats produce diverse types of sounds, and hence members of the genus Felis also growl, hiss, snort, and produce a peculiar snarl. They mainly eat while sitting hunched on the hind legs. The front 923 legs and forearms do not touch the ground. Pieces of the prey are usually bitten off and they do not tear it with upward jerks of the head (see Fig. 32). The geographic distribution of the genus corresponds to that of the family but small cats are evidently absent in some regions within the range of the family. In some countries (Europe and North America) the range has undergone great reduction as a result of extinction during the last century. As pointed out earlier (see characteristics of the family), the genus Felis s.l. represents a progressive group of cats, the members of which reflect most distinctly the main line of specialization of the family. It is the highest type of cat, and it may be, of predators in general. It is closely related to big cats, Panthera. The snow leopard, genus Uncia, is a direct link between Felis and Panthera and a transitional form in the classification adopted in this work. Nevertheless genus Felis is very diverse and includes forms specialized in different directions. Species large in size and adapted to catching relatively big animals belong to this group. While preserving the basic diagnostic features of the genus, these cats include extreme forms which also exhibit a similarity of features with big cats (Panthera) and the ounce. Members of genus Felis include forms transitional between highly specialized species of small felines (sand cat and manul) and ounce and thus big cats (more precisely, the reverse). Examples are the puma and clouded leopard. As mentioned several times before, there is no unanimity of opinion with regard to the composition of the group of cats considered here under the single genus Felis. Some authors divide this group into 12 (Haltenorth, 1957), 14 (Pocock, 1917), 16 (Weigel, 1961) and even 17 (Mazak, 1965) genera. Diversity, undoubtedly considerable, cannot be ignored by adopting a single large genus. This simply emphasizes the gradual evolution of characteristics in the group and the mutual link between what at first glance are dissimilar species or even polar opposites (clouded leopard —sand cat and manul) through a series of transitional forms. At the same time such a division naturally leads to the recognition of subgenera—a category particularly suited for these situations. Various Soviet species have been grouped into three subgenera in this work. The subgenus Prionailurus usually includes three South Asian species which judging from the fourth species inhabiting the Soviet Union (F. euptilura), cannot be supported.* In the subgenus Felis itself different zoologists include five or six Euro-Asian and African species, including the sand cat from Turkestan, the Arabian Peninsula and North Africa which is treated as two separate species (margarita and thinobia). This view cannot *The Amur forest cat produces fertile hybrids with the domestic cat in the Ussuri region (see below). 248 324 be accepted. First of all, these are simply two forms of the same species (Heptner and Dement’ev, 1937), and secondly they differ from true Felis s. str. in several characters, of which the most important peculiarity is the structure of the tympanic bulla. The South African F. nigripes (subgenus Microfelis) should evidently be excluded from this subgenus. The subgenus evidently comprises seven species but its composition differs from that generally recognized: three species invariably regarded as Felis $. str.—silvestris (including group libyca), chaus and bieti and three species usually placed with Prionailurus—viverrinus, bengalensis, and rubiginosus. F. euptilura usually placed in subspecies bengalensis, also belongs here but is treated as an independent species in the present work. Of these seven species, three inhabit the Soviet Union. Subgenus Lynx includes three species of which the Soviet fauna has two and subgenus Otocolobus, two species, both represented in the Soviet fauna. Outside the Soviet Union small groups (two or three species) belonging to fairly distinct or identifiable subgenera exist. More importantly, some individual species deserve elevation to the rank of subgenus (ocelot, serval, flat-headed cat, F. planiceps, jaguarundi, clouded leopard puma, and others). Applying the same principles as used for evaluating relations between genera in the family relations between the subgenera and individual species of the Soviet (and in part American) fauna listed above are as follows. Of the three subgenera, two (Felis and Otocolobus) may be considered forms oF. manul Subgen Otocolobus oF. margarita F. rufao | group о group /ibyca © canadensis Е. ° Subgen F. lynx silvestris / о / eta vex ори ое Subgen Felis Е. caracalo——————— oF. chaus oF. euptilura Fig. 155. Systematic relations between some cats of the genus Felis in the northern Holarctic. V.G. Heptner. 249 325 representing one line (“‘direct’’) of development (specialization) toward the attainment of a completely feline type. Subgenus Felis, however, is less specialized than Otocolobus. The third subgenus, Lynx, represents a lateral line, highly similar and parallel to the general line of the first two, but not attaining the level of specialization of Otocolobus insofar as skull structure is concerned. The main feature of subgenus Lynx is the general body build, which is very typical and exhibits no variation in the various species of this subgenus. These cats stand on long legs and sport short tails; the contour of the body and legs therefore approaches the form of a square. The subgenera Felis and Otocolobus are characterized by long bodies, long legs, and a relatively long tail. The progressive specialization of the skull in these two subgenera has reached (in several species) a level far higher than in subgenus Lynx, and the highest level in the family (subgenus Otocolobus —О. manul). Insofar as relations between species are concerned in the main line (Felis* —Otocolobus) these are defined by a progressive reduction in the facial region of the skull (and a corresponding increase in the cranial region), a progressive turning of the orbits forward, and a partial alteration in the structure of the auditory apparatus. Highly specific adaptations have developed against this ‘*phyletic’’ background (for details, see ‘‘Description’’ of the species and particularly the section ‘‘Systematic Position’’. On the whole, our fauna is represented by the following series: F. (F.) euptilura (Amur cat)—F. (Е.) chaus (jungle cat) —F. (F.) silvestris, group silvestris (European forest cat)—F. (F.) silvestris, group libyca (steppe cat)—F. (O.) margarita (sand cat)—F. (O.) manul (manul) (Fig. 155). Species of subgenus Lynx (lynx), which exhibit the main features of the genus maximally, 1.е., structure of body and legs and general degree of specialization of the skull (in the direction Felis—Otocolobus), form the series: F. (L.) caracal (caracal)—F. (L.) lynx, group [ух (Eurasian lynx) — F. (L.) lynx, group canadensis (American lynx)—F. (L.) rufa (bay lynx [bobcat] of America). Specialization (progressive advancement) of the skull in this series of forms (species and subspecies) and differences in the skull features between the initial and terminal species in the group (caracal— rufa) are less than in members of the corresponding main series (Felis — Otocolobus). The relative level of specialization among species of the subgenus Lynx (lynx), compared to that of the main line, is such that the skull of F. (L.) caracal (caracal) corresponds, or almost so, to the skull of F. (F.) chaus (jungle cat), while the skulls of the two groups of true lynxes, F. (L.) /ynx *Felix in Russian original —Sci. Ed. 326 (lynx and canadensis) and Е. (L.) rufa (bay lynx), reach only the level of Е. (F.) silvestris, group libyca, or slightly higher, but not the level of F. (O.) margarita (sand cat) (Fig. 155). Paleontological data pertaining to the genus are extremely meager. The genus Felis, like the genus Panthera, obviously evolved in the Pliocene and its differentiation is of recent origin. Moreover, a true lynx, F. (L.) brevirostris has been found in the upper Pliocene. This group originated earlier than the Early Pleistocene (Villafranchian)—F. (L.) issidoriensis, through F. (L.) teilhardi to present-day lynx, appearing in the Upper Pleistocene. A distinct species of puma, F. (P.) inexpectatus, is known in the Pleistocene of America (Thenius and Hofer, 1961). The genus Felis is very rich in species of carnivores, in fact one of the richest genera among mammals in general. It contains 29 extant species*: Е. bengalensis Kerr, 1792—A; Е. rubiginosus Geoffroy, 1834—А; Е. viverrinus Bennett, 1833—А; Е. euptilura Elliot, 1871—A; Е. planiceps Vigors and Horsfield, 1828— А; Е. marmorata Martin, 1836—A; F. badia Gray, 1874—A; F. temminckii Vigors and Horsfield—A; F. aurata Temminck, 1827—А; Е. pardalis Linne, 1766—SA; Е. wiedi Schinz, 1821—SA; РЕ. pardalis Linne, 1766—SA; F. иле Schinz, 1821—SA; F. colocolo Molina, 1782—5А; Е. geoffroyi d’Orbigny and Gervais, 1843—$А; Е. pardinoides Gray, 1867 (tigrina) — SA; Е. guigna Molina, 1810—SA; Е. pajeros Desmarest, 1816—5А; Е. yaguarundi Lacepéde, 1809—5А, CA and МА; Е. chaus Giildenstaedt, 1776— А and Af; F. silvestris Schreber, 1777—A and Е (including Е. libyca Forster, 1780— АЕ A, and Е); F. Мей Milne-Edwards, 1892—А; F. serval Schreber, 1776—Af; Е. lynx Linnaeus, 1758—A, Е, and МА; Е. rufus Schreber, 1777—МА; Е. caracal Schreber, 1776—А and Af; F. nigripes Burchell, 1882—Af; F. margarita Loche, 1858—A and Af; Е. manul Pallas, 1776—А; Е. concolor Linne, 1771—SA, CA, and Ма; F. nebulosa Griffith, 1821—A.5 Species of this genus comprise 82% of the total number of species of the family. Most (17) are from Asia; moreover all, except one (lynx), are distributed in the southern half of the mainland. Of these, four are common to Africa, two to Europe, one to Europe and North America, and twelve “This list is only a catalog and does not represent a specialization sequence. E = Europe, A = Asia, Af = Africa, NA = North America, SA = South America, CA = Central America. ‘This list does not include Mayailurus iriomotensis Imaizumi, described in 1967 as a new genus and species from шото Island, one of the southernmost islands of the Ryukyu chain lying on the latitude of Taiwan. This form is described by the author as coming close to the South American F. guigna (‘‘Oncifelis guigna’’). The absence of specimens and the nonavailability of a full description of the species do not permit a positive judgment about the position of this cat from Iriomote. It is quite clear, however, that firstly, another new - genus has no justification from the viewpoint adopted here and, secondly, this cat probably belongs to the group Prionailurus (it greatly resembles F. viverrina and partly Е. еир ига). 250 327 solely Asiatic. In South America there are nine species, of which two are common to Central and North America and seven exclusively confined to the mainland. In number of species Africa stands third with six, of which four are common to Asia or Asia and Europe, and only two are endemic. In North America there are four species, of which one is common to Eurasia, two to South and Central America, and only one endemic. In Europe there are two species, of which one is common to Asia and North America and one to Asia and Africa. There are no endemic species in Europe. Evidently there are two clearly distinct centers of cladogenesis of cats—South America and Asia (southern half). If one takes into account the respective number of species of big cats (Panthera) and ounce, the link between most species of the family from southern Asia becomes even more striking. The center of development of present-day species of the family would seem to lie in this part of the globe, probably in the extreme south and southeast sectors. These are fur-bearing animals of little economic value. Some species are large (lynx, but more so puma) and objects of sport. They control rodent and hare populations and sometimes even large game. Occasionally they harm game populations, primarily through decimation of game birds, and sometimes domestic horned animals. Six species are represented in the fauna of the USSR: Amur cat, К. (Felis) euptilura Elliot, 1871; European wild and steppe cat, F. (Felis) silvestris Schreber, 1777; jungle cat, Р. (Felis) chaus Gild., 1776, caracal, Е. (Lynx) caracal Schreber, 1776; lynx, F. (Lynx) lynx Linn., 1758; sand cat, Е. (Otocolobus) margarita Loche, 1858; and manul, F. (Otocolobus) manul Pall., 1776. These species constitute about 21% of the species of the genus and about 8.0% of the total number of mammals in the Soviet fauna. The range of the genus Felis in the Soviet Union covers all of the forest zone, including montane forests in the southern part of the country, the semidesert zone (in part steppes), and desert regions. The range has shrunk greatly at places in the west in recent years. In certain areas (Middle Asia) they are fur-bearing animals of no mean consequence. In other regions in the south they hurt commercial hunting by decimating game birds. Some species are of vital importance in the live animal trade. (V.H.) oil 328 AMUR, OR FAR EASTERN FOREST САТ® Felis (Felis) euptilura Elliot, 18717 1792. Felis bengalensis. Kerr. Animal Kingdom, p. 151. Southern Bengal. 1837. Felis chinensis. Gray. Mag. Nat. Hist., vol. 1, p. 577. China. Fixed as Canton (Allen, 1938; Ellerman and Morrison-Scott, 1961 and 1966). 1862. Felis undata. Radde. Reisen Stiden Ost-Sibirien, vol. 1, р. 106. Amur, 60 km below mouth of Zeya. Nec Felis undata Desmarest, 1816. 1871. Felis euptilura. Elliot. Proc. Zool. Soc. London, Dec. 1871, p. 761. Substitute for undata Radde. 1872. Felis microtis. Milne-Edwards. Rech. Hist. Nat. Mamm., p. 221. Near Peking. | 1872. Felis decolorata. Milne-Edwards. Ibid., р. 223. Near Peking. 1904. Oncoides bengalensis Kerr a. raddei. Trouessart. Catal. Mamm. Quing., Suppl., p. 271. Replacement for undata Radde, 1862. 1922. Felis manchurica. Mori. Ann. Mag. Nat. Hist., vol. 10, p. 609. Near Mukden. 1929. Felis viverrina mandjurica. Baikov. Okhotnik, vol. 4, р. 16. Northeastern China (Manchuria). (V.H.) Diagnosis Body spotted throughout but spots on back may be faint; color of spots rusty. White field present on back of ear. Two light-colored stripes run from inner corner of eyes onto forehead. Skull elongated; plate in interpterygoid region extended posteriorly (Fig. 158**); and interpterygoid vacuity very narrow. п the most recent period, as a result of assigning the name bengalensis to this species, it is sometimes referred to as Bengal cat. Such a modification of the Russian name, which came into common usage long ago (Satunin, 1914; Ognev, 1935; Novikov, 1956; Stroganov, 1962; and others) is unnecessary and only leads to confusion. ’The species name euptilura is assigned with some reservation. It is possible that the name microtis has some completely formal advantage over it. According to some data the work of Milne-Edwards published between 1868 and 1874 should be regarded as belonging to 1871, especially the earlier sections (Trouessart, 1905; April— Sowerby, 1923), but Elliot's work was published in December, 1871.* According to Ellerman and Morrison-Scott (1951 and 1966), however, the name microtis is dated 1872. Finally, sometimes the names assigned by Milne-Edwards in the above work are assigned simply to 1874. In any case, continuation of the name euptilura is desirable as a nomen conservandum. There is sufficient justification for concluding that the form chinensis belongs to another species (bengalensis, Heptner, 1971). If, however, it were proven that it was conspecific with the Amur form, it is clear that the name chinensis Gray, 1837 should have distinct priority as a species name. *1971 in Russian original — Sci. Ed. **Misprinted 156 in Russian original — Sci. Ed. `лозешоу “n'y Aq Sunureg lla 24 udna (5124) stjaq Yeo лигу © Ald 329 251 252 Fig. 156. Amur cat, Felis (Felis) euptilura (yearling female. ‘‘Kedrovaya Pad’ ** preserve near Vladivostok. October 27, 1965. Photograph by А.С. Pankrat’ev. Second premolar (first in row) usually present; antero-inner cusp of upper carnassial tooth weakly developed and without prominent cusps. (V.H.) Description Small animal, roughly the size of Caucasian forest cat [Felis silvaticus caucasicus}. In general build and appearance this species is similar to the Caucasian forest cat but stands slightly taller on its legs, though not as tall as the jungle cat, and is lighter (Fig. 156*). Tail somewhat shorter than one-half of body length, but usually more than double the length of hind limb. Head small, fairly narrow at the zygoma, and hence facial region pointed and somewhat extended. Ears quite large, close-set, and project prominently from pelage, even in winter; tip of ear somewhat blunt and devoid of tuft of long hair. Hair on cheeks not long (side-whiskers absent). Eyes small with a vertical pupil. Pelage throughout body of uniform length; tail furry and appears quite thick. **Pelage similar to that of European forest cat. On the back an average of 7,000 hair occur per cm? and on the abdomen, 1,950. There are 30 *Misprinted 157 in Russian original —Sci. Ed. N 330 Fig. 157. Head of an old male Amur cat, Felis (Felis) euptilura. *‘Kedrovaya Pad’ *’ preserve near Vladivostok. October 3, 1965. Photograph by A.G. Pankrat’ev. underfur for every top hair. Pelage on back consists of six categories of hair, and five on the abdomen (category IV top hair absent). Average length of guard and top hair on back 47, 49, 47, 39, and 35 mm, and their thickness 108, 106, 75, 66, and 34 microns; corresponding values for hair on the abdomen 45, 43, 37, and 31 mm, and 60, 56, 40, and 41 microns; length and thickness of underfur on the back 34 mm and 18 microns, and on the abdomen 28 mm and 16 microns. Thus there is very little difference in the length of hair on the upper and lower surfaces of the body. The shortness and thickness of guard hair and the absence of one category of top hair on the abdomen are characteristic features’’ (B.F. Tserevitinov). Typical coloration spotty but extremely variable. In extreme cases the entire body of one animal was covered with spots, while spottedness was almost absent in another, standing out prominently only in a small area. The color difference between extreme variants is so great that it has provided 331 а basis for some 7001051515 to propose the existence of two types of small forest cats—one in the Ussuri territory and the other in northeastern China (Maak, 1861; Baikov, 1915 and 1929). The spotted type of coloration, when fully developed, is as follows: Background color on back and upper part of flanks in winter coat usually variable—dirty grayish-pale-yellow or dull grayish-brown with a slight rusty or reddish hue. Color gradually fades downward on flanks, becoming whitish ocherous. Abdomen, groins, and inner sides of legs very light-colored, ocherous or dirty white, or white with a light ocherous dusting. Chin, throat, underside of neck, and chest have a white background. Throughout the body, extremities, along the neck, and in part along the head, occur small (1.0 to 2.0 cm) complete spots of circular or slightly elongated shape; the latter are particularly evident on the back. They have in part sharp outlines, and in part their margins are blurred and indistinct. They are either rusty-ocherous or rusty-cinnamon, light and bright, or duller and fuller. They may be very close-set; sometimes the main background color occupies almost the same area as the total area of the spots. The spots are denser and their total area relatively greater in this species than in other Soviet species of small cats. Three or four narrow stripes extend along the spine consisting of elongated, narrow, sharp, and richly colored small spots. Sometimes their outlines are so indistinct that a generally brown, indistinct, broad belt is formed. Spots and longitudinal rows of sharply outlined spots stand out prominently only at places, mainly on the shoulders and sacrum. Spots on the abdomen and chest are more rounded than on the back, bright, and rusty or rusty-brown in color. Down the throat and neck, the spots are bright rusty or a much darker rusty tone. They are usually disposed in such a way as to form three to five transverse rows, but often fuse into continuous transverse stripes. Rows of spots, three to five, fusing or almost fusing into stripes, sometimes occur across the upper part of the forelegs. Spots on the ‘extremities become increasingly smaller and extend to the tarsal and carpal region. Vertical stripes or vertical rows of spots in a pattern do not occur. The tail is dark gray, turning somewhat lighter in color toward the lower part, and is either monochromatic or with fairly distinct dark (blackish- gray) transverse broad stripes or bands. Tip pure black or dark gray. Throughout the tail an extremely faint ocherous dusting is sometimes present and the stripes thus may have an admixture of rusty tones. As many as seven stripes may occur on the tail. Coloration of head also complex (Fig. 157*). Upper lip, area around eyes, as well as lower lip white, but rust-colored spots occur in the region *Misprinted 158 in Russian original —Sci. Ed. 532 ; ‹ я Мел Ах NX: 4 AY КА ae _ sf - Е ВО pe). М, Ир ет › Se Te К о ‘ИЩИ: 255 Fig. 158. Skull of Amur cat, Felis (Felis) euptilura. №. $ 14197, collection of the Zoological Museum, Moscow University. Cape Gamov near Vladivostok. February 12, 1927. Sketch by N.N. Kondakov. of the whiskers. Upper part of nose grayish with a rusty dusting. From the nose through the forehead to top of head there occurs a broad, dark, reddish- 383 cinnamon stripe, set off on both sides by two distinctly prominent narrow, light-colored (white) fields. This particular marking commences at the inner corner of the eye and extends through the forehead. Beyond, the stripe and white fields diminish and disappear in the general dark color on top of the head. The light-colored fields are bordered by a narrow dark field, somewhat darker than the space between the eye and the ear and the entire upper part of the head. Down the cheeks and behind the eyes a light-colored field occurs, along which two bright stripes of different intensities of reddish, rusty, or light brown color extend posteriorly from the region of the eye. This particular marking merges into the pattern on the throat and the lower side of the neck, or touches it in some manner. On the dark-colored sinciput, occiput, and upward on the neck, sometimes almost up to the shoulder, four narrow brown or rusty-brown fields occur. The back of the ear is, for the most part, white or almost white; just below the tip it is blackish-brown; the tip itself is light rusty in color. The second type of coloration, when fully developed, is as follows. Back and upper flanks indistinct, dirty, brownish gray, with an admixture of silver at places due to the presence of many light-colored top hairs. Color richer and darker on the back, and somewhat lighter on the flanks, especially ventrally. Spots not visible against this main background. Body essentially monochromatic, brownish-gray. Spots faintly discernible along undersides of flanks but distinct on abdomen, where their brown shade stands out prominently against the ocherous white field. Spots indistinct on legs. Spots distinct against general background of light-colored body; nevertheless they are rather vague, diffused, and rusty in tone. Some spots on abdomen and thighs have a small light-colored center. Tail dark, dirty gray, with darker transverse spots. Reddish-colored spots are absent throughout the body in this color pattern; instead, spots are brownish-rusty on the throat, down the neck, and on the groin. The head is a dark or coffee-brown color and the stripes along the sinciput, occiput, and up the neck, dark brown or even blackish-brown. Very distinct light- colored fields extend from the eyes to the forehead. The two extreme types of color patterns described above are often blended in various ways. The main background may vary from dark gray or dark brownish-gray to almost dirty black along the spine (up to the “‘belt’’), to light reddish-ocher and pale yellowish-ocher. Spots may be close-set through the back, flanks, and down the body, or sparse, or not distinguishable, or only barely discernible on the spine; sometimes spots are more “‘blotches’’ than spots. In dark individuals spots may be totally absent down the middle of the back but quite distinct on the flanks, or absent throughout the body except for the abdomen. Sometimes spotting is represent- 254 334 ed by faint shading. Rarely, spots on the thighs and abdomen will have a small, indistinctly contoured, lighter-colored area in the center. Spots may vary from vivid red to dark brown, without entirely “‘reddish’’ [intermediate tones] (even on the abdomen) and dirty black. Stripes on the occiput and neck may be distinct, black, and long, reaching up to the shoulder, or almost imperceptible. Corresponding to the general color tone and intensity of color of the coat, the color pattern on the upper part of the head also varies. Stripes on the cheeks may be bright reddish or rusty, or dark and brown, but invariably exhibit a distinct rusty tone. Spots and transverse bands on the throat vary from bright ocherous-rusty to dark brown with a rusty hue. The tail likewise varies in color, from an almost monochromatic gray in dark- colored animals to gray with faint spots, even though the body spots in some animals may be very vague. Judging from available, albeit meager ав, different types of variegated color patterns are found in the population, while dark, monochromatic animals are quite rare. According to some authors (A.G. Pankrat’ev) the coat may fade significantly after winter. However, both spotted and mono- chromatic cats are encountered in early winter. On the whole, color variation in the Amur cat is greater than that of any other species of Soviet cat, except lynx. However, in all color types the light-colored stripes extending from the corner of the eye and the reddish hue of the pattern on the throat and down the neck remain unchanged. The rusty shade in the fields along the cheeks is also constant, as is the color of the back of the ear.® Not much is known about the summer coat. It is evidently characterized by the presence of sharply visible spots, but the overall range of variation characteristic of the winter coat is also seen in summer. Sexual dimorphism is absent and age-related changes have not been described for Soviet forms. In any case, by the first winter young kittens have developed an adult coat. Geographic color variation throughout the species’ range under consideration here is not adequately known but seems to be insignificant. The skull is notably elongated, with an extended braincase and close- set zygomatic arches, and is low. Upper line of profile bulges slightly and uniformly without prominent elevations in the interorbital region. Frontal region small, narrow, gently bulging, and median longitudinal groove absent. Supraorbital processes well developed, long, usually approaching postorbital “These descriptions of coloration of Amur cat vary greatly from those given in Soviet literature, including even those given for purposes of identification. In the latter a rich and vivid reddish spottiness throughout the body is invariably emphasized as the main characteristic. The present description is based on data from the Zoological Museum, Moscow University. but mainly on new data furnished by A.G. Pankrat’ev (Vladivostok). 256 835 processes of the jugal, but sometimes (more often in old animals) fused, in which case orbit is closed posteriorly. Sagittal crest in posterior part of parietals distinct but low; lambdoidal crests well developed and more complete than sagittal. Rostral part of skull relatively short and broad. Orbits oval, significantly extended backward, and their ventro-anterior region greatly thickened. Maximum diameter of infraorbital foramina wider than space between these openings and edge of orbits. Nasals narrow in posterior half but extended and broad in anterior part. Hard palate narrow and long—its posterior margin lies in interpterygoid vacuity beyond the line joining posterior surface of molars, at a distance more than one-half width of interpterygoid vacuity (see Fig. 158). Interpterygoid vacuity narrow. Choanal openings small; diameter only slightly more than height. Tympanic bulla large, greatly inflated, and long; anterior margin lies close to postglenoid process but does not reach its level. Ectotympanic chamber small but its boundary with entotympanic chamber not visible externally. Distance between inner margins of tympanic bullae equal, or almost equal, to width of interpterygoid vacuity. Canines thin, sharp, and moderate in length. Second (first in row) upper premolar usually present but may be shed during lifetime and even its cavity filled in most adults, especially older animals. Both teeth absent in 30 to 40% of animals examined (A.G. Pankrat’ev). Antero-inner cusp of upper carnassial negligible, or weakly developed, and without sharp edge. Sexual dimorphism in skull absent apart from somewhat smaller size of female skull. Skull of young animals not as long but with larger cranium. Even in the first winter, differences between the skull of young animals and that of an adult diminish notably. Geographic variation of skull mainly concerns overall size, which can be rather significant. For animal measurements see ‘‘Сеостарыс Variation’. (V.H.) Systematic Position The Amur forest cat was initially (Radde, 1862) placed under Felis undata Desm., a species of the Malay Archipelago. Later it was clarified that the Amur forest cat is not related to this species and hence it was (by means of a substitute name) treated as a distinct new species, Felis euptilura (Elliot, 1871). In the various nomenclatural schemes adopted in Soviet and foreign literature, the Amur forest cat was treated as a distinct species of forest cat found in the Amur-Ussuri territory and northeastern and eastern China until the early 1900’s, when it was suggested that the entire group of forms in the species euptilura be regarded simply as a subspecies of the south Asian species, Е. bengalensis Kerr, 1792 (Trouessart, 1904). This suggestion was accepted by several prominent zoologists (Pocock, 1939; Ellerman and 336 Morrison-Scott, 1951; and others), albeit rather tardily. In foreign literature this was generally accepted. Soviet zoologists, on the other hand, clung to the independence of the species Е. euptilura (Ognev, 1935; Bobrinskii et al., 1944; Novikov, 1956); however quite recently, a few have now begun to favor the view that the Amur cat should be assigned to the species bengalensis (Bobrinskii, Kuznetsov, and Kuzyakin,1965). A special study of this problem (Heptner, 1971) has revealed that the Amur cat is more correctly isolated as an independent species, differing significantly from F. bengalensis of southern Asia. The Amur cat is characterized by a number of peculiarities, which distinguish it not only quantitatively but also qualitatively. Compared with F. euptilura, the skull of F. bengalensis has a relatively larger cranium, the lyriform figure on it is larger, and the lines forming it, in fully grown animals, converge not in the region of the frontals or the coronal suture, but farther behind, roughly at the middle of the parietals. The sagittal crest is not developed and appears only at the extreme posterior end. Compared with the skull of the Amur cat, therefore, the skull of F. bengalensis is slightly more juvenile—the skull of adults resembles that of younger Amur animals. In this respect, as shown above (see characters of the family) the skull of F. bengalensis does exhibit some ‘‘progressive’’ features. In these it is similar to the skull of cats of the F. silvestris type. The skull of the Amur cat exhibits structural features which are transitional between those of F. bengalensis and F. viverrina and, in some respects, are closer to the skull of the latter species. However, it is significantly smaller and the crests are less developed. Of greater significance are the color differences between Amur cat and F. bengalensis. Coloration and spottedness are completely different in the Bengal cat. Spots in the latter are invariably sharp against a very light- colored background and readily differentiated in form and size. Small spots occur with very large spots, mostly irregular in form, but often elongated, especially on the spine and close to it. Tail coloration also differs. In the Amur cat some individuals are almost monochromatic; dark or very faintly spotted patterns are normal. If, however, the spots are distinct they are numerous, identical, small, densely set, and do not tend toward elongation. This is the more prevalent type of color pattern (see ‘“Description’’). This type of spotted pattern (leaving aside the relation of spots and background) is similar to that in the steppe cat (F. silvestris caudata). In general, however, the coloration, in particular in the sharply spotted type found in the Amur cat, quite closely resembles the coloration of F. viverrina.? Color characteristics, of vital importance among cats, reveal a greater affinity with °See sketches of F. bengalensis and Е. viverrina presented by В.Т. Pocock (1939) and coloration descriptions of species F. bengalensis and Е. euptilura given by V.G. Heptner (1971). 257 337 Е. viverrina than in skull structure. On the whole, while belonging to the same group, ‘‘Prionailurus,’’ as do Е. viverrina, Е. bengalensis, and Е. rubiginosa (possibly also F. planiceps), Е. euptilura occupies an intermediate position between F. viverrina and F. bengalensis. Moreover, the delimitation of this group, which at times is considered a subgenus of genus Felis and at times an independent genus, is not well defined; neither is its compactness. Thus some workers break this ‘‘genus’’ into several separate subgenera distinct from genus Felis—viverrina (Zibethailurus) and planiceps (Ictailurus). Moreover, the entire group **Prionailurus”’ exhibits a very close relationship with the conventional group Felis s. str. and, as mentioned earlier, its further classification into species is difficult to substantiate. Hence, here, F. euptilura is placed in genus Felis $. 1. and subgenus Felis. In skull structure (see characters of the family and genera Panthera and Felis), Е. euptilura occupies, among Soviet species of the genus, the position of least specialized or one of the least specialized forms. This species stands at the beginning of the series which leads to the more progressive conditions culminating in development of the “‘feline’’ type in genus Felis $. 1.— the manul; it is closest to F. chaus (jungle cat). The proximity of Amur cat to species of Felis s. str. and the incorrectness of separating it as an independent subgenus, or even a genus, are supported by the fact that the Amur cat hybridizes with the domestic cat in natural conditions. Crossbreeding often takes place in forest villages in the Ussuri, the male partner being the wild cat. In a backcrossing of hybrid males and females with domestic males and females, progeny have been obtained which are fertile in both combinations. Furthermore the hybrid generation is fertile among its own members (A.G. Pankrat’ev). For body measurements, see “‘Geographic Variation’’. (V.H.) Geographic Distribution The Amur cat inhabits forested regions of the Amur—Ussuri territory and northeastern and eastern China. Geographic Range in the Soviet Union The range in the Soviet Union (Fig. 159) represents the northeastern boundary of the range of the species. It encompasses the southern part of the Far East and associated forests of the so-called Manchurian type. The range is interrupted within the Soviet Union but the two sections merge outside the boundaries of our country. In the west the range boundary, commencing at Blagoveshchensk on the Amur, runs toward the middle course of the Tom’, the left tributary of 338 <, BER RRS owas XS 6% Pats QO А 80 160 240km Fig. 159. Distribution of Amur cat, Felis (Felis) euptilura, in the Soviet Union (scale in km). 1—northern boundary of range according to data up to the early 1930's; 2—same, up to 1968; 3—discontinuous (diffuse) distribution; 4— dense distribution and region of highest population density in 1968; 5 — possible habitation up to Gorin mouth in the past. V.G. Heptner (distribution and population density in 1968 according to data of А.С. Pankrat’ev and С.Е. Bromlei). the lower Zeya, roughly toward 51°17’ N. lat. (northernmost point of 258 occurrence). Turning east and slightly south it then intersects the Turan range and the middle course of the Bureya, runs toward the upper Mutana, and descending from there to 49° N. lat., turns toward the upper Bidzhan. From there it descends southward and enters China. On the right bank of the Amur, the northern boundary of the range is formed by the Amur in the stretch from the state boundary and the mouth 339 of the Ussuri in the west to the mouth of the Sungari in the east. From this point (50°06' М. lat.) the boundary runs southeast and south along the sources of the Monoma, Anyuy, and Khor, descending to the Mukhen’ River around 48°55’ N. lat. From there it runs eastward, intersects the upper Khor, and through the Murav’ev-Amur pass to the Samarga River and the sea of Japan at 47°20’ М. lat. In the expanse between Ussuri and the ocean, the Amur cat is confined to nut pine—broad-leaved, and mixed forests, and does not ascend beyond 500 to 550 m above sea level. In other words this species is absent in the higher altitudes of the Sikhote-Alin covered by Okhotsk taiga. It is possible that, in the north, it formerly reached the mouth of the Gorin (50°45’ М. lat.). This species breeds on Russkii Island at Vladivostok. The boundary outlined above, although schematic (range in Ussuri territory around Sikhote-Alin), summarizes data available on the distribution of Amur cats from the time of the first Russian explorers of the Far East (1850’s) to the early 1930’s (Ognev, 1935; Zubarovski, 1939). The 1968 boundary was probably of the following form. Commencing in the west slightly east of Blagoveshchensk on the Amur, it ran southeast along the river at distances from it varying from 30 to 50 km, to 131° Е. long. There it sharply deviated from the river, turning initially straight eastward and later southeast, and, intersecting the Amur, entered China. This arcuate boundary covered the entire Bol’shoi Bira basin except the upper portions of its left tributaries, and crossed the Amur at some 40 to 50 km below the mouth of the Bol’shoi Bira. Within the Soviet Union, Amur cats were not found at the time of the survey farther down along the left bank of the Amur River. The boundary again re-entered the Soviet Union at the mouth of the Ussuri but Amur cats were absent in the immediate proximity of Khabarovsk. On the right bank of Amur, the boundary extended to the mouth of the Sungari. There it turned slightly eastward into the mountains, and then turned sharply southward along the western slope of the Sikhote-Alin through the upper Anyuy, Khor, Bikin, and Iman, without embracing, however, their sources and uppermost reaches. The boundary extended in this direction south to 45° or 45°15’ М. lat. The boundary ran here for а distance of 120 to 150 km from the Ussuri. At this point in the south it intersected the main axis of the mountains and turned immediately northeast, running along the eastern slope of the Sikhote-Alin. It continued north in this direction to the mouth of the Botcha (Grossevichi) at 48° N. lat., where it reached the sea. This part of the range is in the form of a very narrow strip at its widest point in the south (about 60 km wide slightly north of Terneya Bay). Thus the extensive, higher parts of the Sikhote-Alin, covered by Okhotsk type taiga or devoid of forests, did not fall within the range. Farther south the range 340 covered the entire territory of the Ussuri krai (from data of А.О. Pankrat’ev and G.F. Bromlei). Boundary differences between the 1930’s and end of the 1960’s are primarily the result of precise demarcation in recent decades (Sikhote-Alin, left bank of Amur). However, some reduction in the range has also been brought about by the extinction of the animal in the north where its population has always been sparse (see subdivision ‘‘Population’”’ under the section ‘‘Biology’’ below), from changes in natural conditions (clearing of forests | | Е =, =] =) all j i | рт М i | 1 | mp | } 4 И 4 Oo | СЕБЕ © EE SS i 0 1000 2000 3000 4000km i Fig. 160. Distribution (scale in km) of Amur forest cat, Felis (Felis) euptilura EIl., and Bengal cat, Felis (Felis) bengalensis Kerr. Lower reaches ог Yangtze — southernmost point from which Amur cat is known (Nanking; Heptner, 1971). Southern boundary of range of Amur cat calculated as 30°N. lat. V.G. Heptner. 259) 341 and shrubs). Adverse effects in one part of the range are naturally reflected in other parts. Evidently boundary variations, although not directly proved, also arise from natural climatic and landscape (forests of Manchurian type) limits to the range. Existing under such conditions the species must have become sensitive not only to long-term climatic changes (they could have remained adequate for over a hundred years) but also to temporary fluctuations. Geographic Range outside the Soviet Union How large the size of the range was outside the Soviet Union (Fig. 160) (related to the question of forms) cannot be stated with certitude since the species limits are not yet clear (see “‘Systematics’’ and “‘Geographic Variation’’). In any case, it includes northeastern China (evidently only the eastern and northeastern parts), Korean Peninsula, the islands of Cheju Do and Saisyu (Tsushima and Quelpart), and eastern China, at least to the lower course of the Yangtze Kiang (Nanking"®). It is possible that this cat occurs even farther south but it is not known precisely to what extent and where the boundary lay in the past in relation to that of F. bengalensis, nor whether these two species are (were) encountered together in southern China. In any case, F. b. bengalensis has been recorded from Yunnan; the form described under the name F. b. chinensis from Fukien, Hunan, Shansi, Hopei, and Szechwan, a small cat with some color features of bengalensis and euptilura, judging from descriptions, pertains to bengalensis (Allen, 1938). The distribution of the species in western China is also not clearly known. True F. bengalensis, with wholly typical features, lives in the Democratic Republic of Vietnam. ok 2: * Cats assigned to the species bengalensis, in addition to the foregoing regions, also inhabit Gansu and Szechwan, Kam, Taiwan (Formosa) and Hainan, Indochina and Malacca, the islands of Sumatra, Java, Bali, Kalimantan (Borneo), Palawan, Calamian, Panay, Negros, and Cebu in the Philippines, Burma, Assam, Sikkim, Bhutan, Nepal, Kumaon, Kashmir, Punjab, the Indian Peninsula almost up to the southern extremity (Palni hills at 10° N. lat.), and southern Baluchistan. This species is absent in Sri Lanka. References to its occurrence in Tibet are erroneous or applicable only to Kam. Although the extent of distribution of the species euptilura, in the sense accepted here, is not wholly clear, it is obvious that true F. bengalensis lives in the Democratic Republic of Vietnam and evidently also in southeastern !0Specimen in collection of the Zoological Museum, Humboldt University (Berlin, German Democratic Republic) (V.G. Heptner). 342 tropical and subtropical China. Thus the range of F. euptilura in relation to that of F. bengalensis is allopatric; however the overlapping of their ranges in the south cannot be ruled out. It is interesting that the range of F. euptilura coincides with the region of occurrence of Manchurian-Chinese relicts. The species itself should probably be considered a member of this relict group. It has been suggested (Pocock, 1939) that Е. bengalensis, i.e., all forms of southern Asia from Kashmir to southern Baluchistan, Indochina, the Malay Archipelago, and southern China, and the group euptilura, regarded here as an independent species, evolved (like tiger) in the north and, later, having colonized southward in east Asia, extended still farther southward. As can be seen from the above account such a viewpoint is difficult to accept. If Е. bengalensis s. str. and Е. euptilura have common roots, then Е. bengalensis should also be very old in origin (typical of south Asian species), like several other forms of the eastern (Indo-Malayan) region, i.e., F. rubiginosa and F. viverrina, which are close to it, as well as F. nebulosa, F. marmorata, F. temminckii, Е. chaus, and others. A reverse situation is possible: a distinct colonization of some progenitors of south Asian cats into the north for F. euptilura. (V.H.) Geographic Variation The species range of the Amur cat, in the accepted interpretation of the scope of the species, is not very large. Nevertheless it extends fairly far south where environmental conditions significantly differ from those prevailing in the Amur-Ussuri territory. Its geographic variation has not been adequately analyzed, apart from the normally occurring range of individual variation. At present it is still difficult to assess positively which among the subspecies assigned traditionally to bengalensis $. 1. actually belong to this species, and which to euptilura. This pertains in particular to the form chinensis. Only one subspecies occurs within the Soviet Union. Amur forest cat, F. (F.) e. euptilura Elliot, 1871 (syn. raddei, manchurica, manjurica; the names undata, bengalensis, chinensis, and microtis were once applied to this form). The ‘‘Description’’ given above pertains to this form and has been developed on material from the Ussuri territory. Size large, representing the largest form of the species. Body length of males 60 to 85 cm, and of females 61 to 77 cm; tail length 23.5 to 44.0 cm and length of ears 4.2 to 5.0 cm. Height at shoulder, 31 to 36 cm. Greatest length of skull in males 96.7 to 115.3 mm, and of females 88.2 to 102.0 mm; condylobasal length of skull in males 89.5 to 105.0 mm, and in females 82.3 to 92.8 mm; zygomatic width in males 60.7 to 74.5, and 343 in females 57.0 to 67.2 mm; interorbital width in males 14.4 to 18.8 пит, and in females 13.1 to 16.3 mm; postorbital width in males 27.0 to 28.8, and in females 26.0 to 28.0 mm; width between infraorbital foramina in males 27.5 to 29.0 mm, and in females 25.7 to 27.3 mm; width above canines in males 24.1 to 28.6 mm, and in females 24.1 to 26.0 mm; and length of upper tooth row (with canine) in males 31.0 to 34.5 mm, and in females 27.8 to 31.0 mm. Weight varies from 2.5 to 7.07 kg; one animal weighed 8.2 kg and another 9.914 kg (exceptionally obese male).!! Much bigger, gigantic individuals are extreme rarities in the Soviet Union. Thus, at Birobidzhan a male was caught (December, 1935) with a body length (from nose to anus) of 107 cm and tail 44 cm, and another male north of Vladivostok (Anuchinsk region; February, 1936) with a body length of 103 cm and tail 42 cm. These animals were measured ‘Чт the flesh’’. The skins of some cats correspond in size to those of small lynx (about 100 cm without tail; Zubarovskii, 1959). Such specimens fall so much out of the normal range that they cannot even be regarded as extreme variants. It is possible that they represent heterotic hybrids of Amur with domestic cat (see ‘‘Systematics’’ above). Measurements of forest cat populations of the southernmost part of the Ussuri region are presented in Tables 5 and 6.1? The extremely high variability in weight of the cats is due not only to sex-related and age-related differences, but seasonal changes. The maximum weight of old males occurs at the beginning of winter. By spring it drops greatly, although sometimes the animals are well fed even in January. Much depends on climatic and other conditions in a given year; cats become very obese when small rodents are plentiful; otherwise, they become extremely emaciated even in January and February (А.С. Pankrat’ev). ‘‘Exceptionally obese’’ males and females were caught on October 25, 1946 and January 2, 1945 with a body length of 66 and 62 cm, weighing 5,700 and 4,319 с (С.Е. Bromlei).!° Weight of heart of adult and subadult males (10) 8.3 to 37.0 5 (М 21.04) and cardiac index 3.49 to 7.89 (M 5.35); corresponding values for adult and ‘1Summarized data include those published by М.А. Baikov (1934), 5.1. Ognev (1935), MI. Zubarovskii (1939), S.U. Stroganov (1962), material furnished by М.К. Vereshchagin (collection of the Zoological Museum, Academy of Sciences) and A.G. Pankrat’ev, and collection of the Zoological Museum, Moscow University; in all, some 50 specimens are represented. !2Tables include specimens of the-Zoological Museum, Moscow University, data furnished by С.Е. Bromlei (Sudzukhin preserve), and material of А.С. Pankrat’ev (vicinity of ‘‘Кедгоуауа Pad’ *° preserve, southwest of Vladivostok). 13 changes with the nutritional plane, and also affects the biologically important character of weight load on the sole (see ‘“Biology’’). ‘зоирЯ [е1э4э9— еш8цо ULISSNY ит USAIS JOU URI, 05° OOL‘T 8 Or 4 OV 8 6 Oel АЯ! 8 O€T 6 005 OC 8 S6¢ 6 065 005 8 059 6 о 1 a ow | (yore 0} 1э4шэоэ( 148пео) so[euiay eoA-]sILJ +5э[еэу зару soem уару ALOFT} 11055(] ay} JO zed илэЦ3п0$ шолу viNgydna °a (+41) °`Ч ‘sed 35эл0у лашу JO 3481эм рие 1481Э № sieved JO 1481эН 1003 риН yysus] пет, 413031 Apog э715 Apog °< э1Е, зэхэрц] 197 си O77 9°67 0°67 0’9Т 9'T9 0°06 8°86 Iewah эцо о} dn OTT oT? 8°97 €9¢ OCI TES T08 0°98 зэ1ешэу J[Npeqns И 8°CC CANS 9°97 vor с 19 8`©6 0`<От зэтеэу уиру OTT CCC L'6¢ ГУС ТЕТ 0°09 668 € L6 won do 6 06 6 IT CCl СТ SV | €Ce CCE | 967 005 | TS? ЭТ | OVI 079 | LOS CH | V8 566 | $06 еэА эио о} dn зэеш предп ~r OOH CO OCT | LTT Ose | OZ СИЕ tLe GLE | €C? 88Т | 9ST Crl |819 O'SOT| O'S6 ЕСТ! L TOL хеш | ини soyeul упру 100} Тетзеилео Jaddn jo Ч18иэ] эшиео пом MOI 41001 Jaddn дю ц18иэ7] эшиео YIM MOI 4100} 1э44п jo Ч13иэ] YIPIM 1314701504 UIPIM 1891470191] рим эцешозА7 Ч13 эт [е5еао|АриоЭ 413 49][ 1$2}еэ10) зэхэри] ALOPIIIN LINSSE 9y} JO зле WieyyNos шол} ‘vénzydna °2 (°.J) Я ‘5 15910} INUIY JO ззиоолизези [[NYS °9 equ COT 346 subadult females (10) 10.6 to 27.3 g (М 18.16) and cardiac index 4.54 to 6.23 (M 5.18). Length of intestines in adult and subadult males (11) 120.0 to 189.0 ст (М 159.5); ratio of same to body length 1: 2.30 to 1:3.0 (М 1: 2.57); corresponding values in adult and subadult females (10) 132.0 to 211.0 cm (М 149.6) and 1:2.36 to 1:2.70 (М 1:2.55) (A.G. Pankrat’ev). F. euptilura is found in the Pri-Amur and Ussuri territory. Outside the Soviet Union these animals occur in northeastern China (Manchuria) and the Korean Peninsula. In the accepted interpretation of species euptilura, and in accordance with the earlier nomenclatural discussion (See рр. 335 to 337), the nominal form inhabits the Soviet Union. In any case, labeling them ‘‘Felis euptilura microtis,’’ as has been the practice, or placed in form chinensis, as suggested recently (Bobrinskii et al., 1965) should not be done (Heptner, 1971). Relations between the Soviet form and several geographically adjacent forms are not clearly understood. Evidently this same large form has been found in northeastern China also (Baikov, 1929) and a similar, but much smaller, form seen in the south.!+ This may be called microtis M.-E. (type loc. Peking). Sometimes it is labeled chinensis Gray (type loc. Canton), which is not correct. The name manchurica Mori is evidently a synonym of euptilura. Either 10 or 11 forms are usually included in species bengalensis outside the Soviet Union (Chasen, 1940; ЕПегтап апа Morrison-Scott, 1966; Weigel, 1961). This number is evidently an exaggeration. Of these forms chinensis (see above p. 342 and Heptner, 1971) possibly belongs to species euptilura. It occurs from Fukien in the east to eastern Tibet and Szechwan in the west, Mongolia (Inner Mongolia—V.H.), and Taiwan in the north [sic] (У.Н.). Biology Population. The range of the Amur cat is associated with Manchurian type broad-leaved forests. In the Bureya basin and the Zeya and its tributary, the Tom’ this cat is very rare, and even rarer in Birobidzhan (almost extinct). On the Zeya-Bureinsk plain (‘‘Amur forest-steppe’’) and in Birobidzhan, '4Measurements reported by Shou (1958)* evidently pertain mainly to young animals: body length (seven animals) 46.0 to 55.5 cm; weight (five animals) 1,750 to 2,875 g; and total skull length (four animals) 87.1 to 98.1 mm. In addition to the foregoing, mention is made of one animal with a body length of 66.0 cm and another weighing 7,200 g (female with a body length of 55.0 cm) (V.H.). *Not in Literature Cited— Sci. Ed. 264 347 however, this cat was common before the war. The reasons for its population reduction are the cultivation of virgin lands, burning and felling of forests and thickets which mainly serve as shelters for the animals, and the plethora of hunters who invaded the remaining small areas of habitat. On being chased by adog, an Amur cat invariably escapes into a tree; here it is easily killed and usually thrown away. This species avoids large dense forests (A.G. Pankrat’ev). In the basin of the lower Ussuri it has become fairly rare in recent years. It is still quite common in the southern half of the Ussuri territory (Pos’etsk, Vladivostok, Shkotovsk, Suchansk, and other regions). In each of these regions in the 1920’s some 100 to 150 cats were caught annually. The maximum density in years of high population in the extreme south of the Ussuri territory, in ‘“Kedrovaya Pad’ ”’ preserve south of the Khasansk region, was two to five animals per 1,000 hectares (A.G. Pankrat’ev). Habitat.’> The Amur cat, like most other cats, is not adapted to living in very snowy regions, i.e., in areas where the snow cover over a 10-day period exceeds 20 to 30 cm, or snowfall is protracted. However, this cat finds optimum conditions of existence in sections where the snow cover is less than 10 cm deep (Formozov, 1946). With a very high weight load on the sole of the foot, equalling 188 © per cm? (С.Е. Bromlei),'° the animal sinks in deeply in loose snow, and hence avoids snow covers of greater depths. Confronted with a snow cover the cat usually moves along crusted portions on streams and brooks, trails of other animals, roads, and rocky sections, from which the snow has been blown off by the wind, or under fallen trees and places cleared of snow by marine surf. Deep snow cover deprives this animal of its chief food, 1.е., rodents, since the predator cannot catch them under snow. A second requisite for the occurrence of this cat in a given region is a fairly abundant supply of mouse-like rodents, Manchurian hares, and pheasants, which it usually hunts. The selection of habitat by this animal is related to these requirements. Thus in the southern Primor’e and Ussuri basin it inhabits sparse Manchurian type broad-leaved forests where these dietary items are more common. The predator occurs less often in these regions in nut pine-broad-leaved forests. It does not ascend mountains beyond 500 m above sea level. It is frequently encountered on creeks draining into river valleys, among thickets of rose willow or sedge, where it hunts for hare. In winter this cat is sometimes confined close to unfrozen springs from which it can catch fish (Zubarovskii, 1939). It rarely appears on the western ‘Habitats of the Amur cat are also characterized by photographs of the biotopes of Amur leopard and Amur tiger; the three species are encountered together almost everywhere. 1бАссог пе to А.С. Pankrat’ev, in adult and subadult males (10 animals), depending on their obesity, the weight load per cm? of sole surface was 48 to 115 g, and that in females (10 animals) 50 to 87 в. Fig. 161. Habitat of ше Amur cat in the lower Ta-Chingou River. Suputin preserve, Ussuri territory. August, 1958. Photograph by У.Е. Prisyazhnyuk. slopes of the Sikhote-Alin (N.V. Rakov) because climatic conditions prevailing there are not favorable. Winter in that region extends from mid- November to March and is severe and frosty. Even at the end of November the atmospheric temperature may drop to —35° С. Winter sets in quickly and snowfall is continuous throughout the season in Sikhote-Alin. Thus few Manchurian animals are seen in this region during the winter. Close to the Sea of Japan, on the eastern slope of Sikhote-Alin, the Amur cat lives mostly along the sea coast. A stretch 5.0 to 6.0 km wide along the sea, devoid of human settlements, contains the maximum concentration of this species. In winter the cat is totally absent in sections with a snow cover 30 to 35 cm deep, i.e., along mountain slopes 500 to 600 m above sea level (G.F. Bromlei). Along the coast this cat is most frequently met with in burned-over forest sections with 40- to 50-year-old Mongolian oak trees and often bushes, hazel shrubs, etc., forming thickets impenetrable to man. Manchurian hares and pheasants are abundant in such thickets. The cat readily inhabits forest glades also, as well as forest belts adjoining cultivated fields, and the shores of marine bays. It does not avoid open meadows and untilled lands. In all the foregoing situations, mice, field voles, Manchurian hares, and pheasants are generally plentiful. Furthermore the snow cover in such places is usually not deep and the snow almost throughout winter rapidly becomes 349 265 i) lon Nn Fig. 162. Herbaceous-shrub growth in the Sukhorechensk range, representing year- э 53 round habitat of the Amur cat. ‘‘Kedrovaya Pad’ °” preserve near Vladivostok. August, 1963. Photograph by A.G. Pankrat’ev. covered with a thin crust, facilitating movement of the cat. The predator is confined year-round everywhere to cliffs and rock slides, where it sets up a lair in summer and protects itself from deep snow in winter. Compared to the number of Amur cats found in the foregoing habitats, only a small population will be found in broad-leaved forests of oak, maple, linden, ash, Manchurian nut, philodendron, and others commonly lacking dense undergrowth. In nut pine-broad-leaved forests at heights of 300 m above sea level the Amur cat is still rarer. Finally, it is absent, as a rule, in the zone of dark coniferous taiga, consisting of spruce and fir, that occupies montane slopes 500 to 600 m above sea level. It usually moves into the taiga only in sum- mer and primarily in years when chipmunks and tree squirrels are abundant (G.F. Bromlei). Thus, in Suputinsk preserve in the Primor’e, noted for its nut pine-broad-leaved forests, this cat appears from time to time only in the snowfree period, and confines itself mainly to old clearings where mouse- like rodents, chipmunks, and Manchurian hares are abundant (Bromlei and Gutnikova, 1955). Consequent to a plentiful snowfall the cat experiences 350 a period of food scarcity and descends into the valleys, coming closer than usual to human dwellings. Only during periods of heavy snow are cases of cat attacks on fowl reported. Its appearance in villages was noted time and again, for example, in the very snowy winter of 1935/1936. This cat could hardly have survived on chickens, however, since dogs begin to bark as soon as the birds squawk (Zubarovskii, 1939; G.F. Bromlei). Food. This cat feeds on ‘*mice and various birds, the nests and young of which are mercilessly ravaged’’; it also attacks hares and young roe deer (Przheval’skii, 1870). In summer the predator seeks out nests of marsh and water birds and eats the fledglings; lizards and snakes are also consumed. In the forest, chipmunks, tree squirrels, and young roe deer fall victim (Zubarovskii, 1939). The list of animals preyed upon by the Amur cat in Kirin province of China is roughly identical (Baikov, 1915; except for roe deer). In years of high populations of mice, voles, and chipmunks, when there is no snow, these rodents constitute the primary captures of the Amur cat. In winter chipmunks hibernate, and catching mice, especially voles, becomes more difficult; hence the cat often resorts to Manchurian hares, tree squirrels, pheasants, and hazel grouse. Rarely, it catches small birds. In years of low populations of mouse-like rodents, the role of birds in the diet of this predator increases. Hazel grouse and hare are the main quarry in winter. The cat consumes spawning fish, which it catches along river banks and, in winter, in unfrozen ponds and brooks. The cat very rarely attacks the young of roe deer, sika, goral, and other ungulates and, when it does, only animals a few days old left unguarded by mother wandering far from them. In years of food abundance these cats become extremely obese. Greatly weak- ened cats are caught in years of low mouse-like rodent populations, especially in winter; the stomachs of such cats are either empty or filled with rotten wood and tree fungi, evidence of the animal’s starvation (G.F. Bromlei). In spite of the comparatively small size of this cat, two instances have been established of its finishing in a single sitting an entire pheasant, including much of the feathers (Bromlei, 1951). Apparently in such cases the cat does: not actually eat some of the flesh but hides it for later consumption. Home Range. The size of individual territories in the spring-summer period is not known. In spring in Sudzukhin preserve it was roughly estimated as 9.0 km?. At the end of winter, when snow is particularly abundant, the cat may cover 3.0 to 3.5 km in a single hunt. Once, in March, a cat was tracked: Гог a distance of 3.5 km. It made its hunting circuit at night and returned to almost the same site from which it left, but entered another hollow in a linden tree, remaining there throughout the day. On this hunt the cat caught three mice and one hazel grouse (G.F. Bromlei). In winter the cat has many shelters in which it rests during the day. 266 267 S51 Fig. 163. Montane gulley in “‘Kedrovaya Pad’’’ preserve. Habitat of Amur cat. September, 1963. Photograph by A.G. Pankrat’ev. Burrows and shelters. ш the Primor’e the den for raising kittens is set up in a crevice among rocks and in a hollow. In June, 1936, in the Pos’et region, two Amur cat kittens were found in a burrow (Zubarovskii, 1939). On the eastern slopes of the Sikhote-Alin this cat lives most often in tree hollows (G.F. Bromlei). In northeastern China kittens have been found in such hollows (Baikov, 1915). In southern China lairs have been discovered in tree hollows and under rocks in thickets (Allen, 1939), Daily activity and behavior. The Amur cat leads a nocturnal life year- round and is seen very rarely in daylight hours because it then rests in its den. It is a skillful climber of trees and is usually caught in them when trying to escape pursuing dogs. Seasonal migrations and transgressions. These aspects have not been much studied, but the Amur cat is known to wander. For example, on the eastern slopes of the Sikhote-Alin, the snow cover is never very thick in the first half of winter. In March, however, so much snow may fall in two or three days that it attains a depth of 50 to 100 cm. Given deep, porous snow cover, cat tracks will not be seen unless a crust has formed; evidently this animal migrates at that time into low-snow areas. 267 Fig. 164. Forest in ``Кедгоуауа Pad’ ”’ preserve near Vladivostok. Habitat of Amur cat. August, 1969. Photograph by G.N. Simkin. In the abundantly snowy winter of 1947/1948, when snow fell in November and remained until March, reaching a depth of 85 cm, these cats disappeared even from those sections of Sudzukhin preserve where they had invariably been found before (G.F. Bromlei). No doubt, in addition to migration, some simply died of starvation. Reproduction. This aspect is poorly known. In the southern Primor’e These animals come into heat in March and live at that time in pairs. Gestation, as in other small cats, requires about 60 days. Kittens appear in the second half of May (Zubarovskii, 1939). In the Ussuri valley kittens have been encountered at the end of April or early May (old calendar; Maak, 1861). On the eastern slopes of the Sikhote-Alin in recent years animals in heat have been sighted throughout the month of March. Kittens appear from the first few days of May (G.F. Bromlei). In Sikhote-Alin preserve a pregnant female was caught on May 7th* with eight very large fetuses (V.D. Shemykin and Yu.A. Salmin). In Kirin province of China, kittens have been seen in May (old calendar) (Baikov, 1915). In the south, in Yunnan province (southwestern China), newborn kittens have been found on February 27th *Year here and below omitted in Russian original —General Editor. 268 393 and April 1st (A.A. Sludskii), and three kittens about two weeks old found in southern China on May 29th. Three small kittens were found on Hainan Island on March 26th (Allen, 1938). In India kittens are seen in May (Pocock, 1939). In the Soviet Union litters usually contain two kittens each and more rarely one or three (Maak, 1861; Zubarovskii, 1939); evidently larger litters are possible. This is supported by the capture of the pregnant female mentioned above. Litters of this cat discovered in Yunnan province consisted of two to four young, and in India, of three or four. Growth, development, and molt. The general color tone of the upper part of the head, neck, and back in a newborn kitten is light brown with _areddish hue. The upper lip is creamy-white. From the upper edge of the nostrils a milk-white stripe extends on each side upward through the forehead. These stripes are bordered by two blackish-brown lines. On the small muzzle and forehead there are consequently two light colored stripes and four blackish-brown ones. Along the upper part of the neck and on the back, blackish-brown spots are longitudinally disposed on the main background color, which fuse into approximately eight parallel stripes. The lower lip, underside of the neck, chest, and abdomen are creamy-white, mottled with longitudinal blackish-brown spots. The lower part of all four paws is pale yellow and devoid of spots. The upper portion of the tail is the same color Fig. 165. Female Amur cat, 10 to 11 months old, menacing photographer. **Kedrovaya Pad’ ^° preserve near Vladivostok. April, 1964. Photograph by А.С. Pankrat’ev. 269 354 as the back but slightly lighter and mottled with longish spots. Distally, the tail coloration becomes a monochromatic light brown. The growth and development of kittens have not been studied. It would seem that the young live with their mother far into the autumn, and possibly even until she comes into estrus again, since the tracks of a female with kittens were encountered on November 27. The footprints of these kittens were almost the same size as those of the mother (G.F. Bromlei). Molt in the Amur cat has not been studied. Enemies, parasites, diseases, mortality, competitors, and population dynamics. No information is available about the enemies of the Amur cat except for one report of cat purportedly torn to bits by an eagle owl [Bubo bubo}. Every cat caught in the spring-summer period, especially June, has been infested with mites [Ixodes persulcatus (infestation index 49.3%), Haemo- physalis japonica douglasi (48.0%), and H. concinna (2.7%). These mites overwinter on cats (up to 40 individuals have been counted in some animals). Five species of fleas have been identified: Chaetopsilla appropinguans (12.5%), C. micado (18.75%), Paraceras flabellum (6.25%), Ceratophyllus tamias (6.25%), and Leptopsylla ostisibirica (56.25%). These fleas were detected year-round. P. flabellum is usually specific to badgers, and is encountered in them from the Primor’e along the mountains as far as the Tien Shan (A.G. Pankrat’ev). In 50% of the cats examined tapeworms were present in the anterior part of the intestine, but their species composition, was not established. Infectious diseases of Amur cats have likewise not been studied. In general, competitors of the cat are fox, lynx, and yellow-throated marten, since the food of all four is quite often identical. Population dynamics have also not been studied. Presumably, popula- tions of this cat would vary significantly in relation to fluctuations in stocks of mouse-like rodents and other animals serving as food. In winters of greater than normal snow some cats die from hunger and resultant diseases. In recent years, with an increase in areas cleared, expanded forest felling, and plantations, the cat population in the southern Primor’e has been increasing and the animal is being seen more often in villages (G.F. Bromlei). Present- day small-scale hunting has no adverse effect on the cat population. Field characteristics. This cat differs from the domestic cat in its much larger size, long legs, and comparatively short tail; tail length is equal to one-half of body length with head. The body color is reddish, mottled with small rust-brown spots. On the head appear two prominent light-colored stripes, running parallel posteriorly along the forehead (see Fig. 165). The pace of walking is similar to that of domestic cats, but strides longer. The average paw size, measured from tracks, is front paw 37 mm x 38 mm and 355 hind paw 36 mm x 30 mm. The cat buries unconsumed prey, for example Manchurian hare or fowl, in the forest and invariably returns to it the following day (G.F. Bromlei). (A.S.) Practical Significance The Amur cat is not very important to the fur industry. Throughout its range in the Soviet Union, in the best years only 1,000 to 2,000 skins, costing 1,000 to 1,500 rubles, were tanned. In the Far East area the following number of skins were tanned: 1,260 in 1932, 2,177 in 1933, 886 in 1934, and 1,301 in 1935. Of the 34 regions in which the skins of this cat are tanned, the most important are Khankaisk, Grodekovsk, Shkotovsk, and Pos’etsk. In each of these regions over 150 skins were tanned in the 1930’s, followed by the Vladivostoksk, Budennovsk and Voroshilovsk regions, in which 50 to 150 skins were tanned in a season. Even smaller numbers, i.e., 10 to 50, were tanned in the OI’ ginsk, Khorol’sk, Chernigovsk, Ivanovsk, Anuchinsk, and Terneisk regions (Zubarovskii, 1939). In the Primor’e territory from 1935 through 1946 the number of cats caught annually was 104 to 269 (G.F. Bromlei). The catch has decreased greatly in recent years, with the total output of skins in the Soviet Union placed at only 100 to 300. By destroying rodents, chipmunks, and hares, the Amur cat often protects crops from pests and hence should be considered a useful animal. It rarely attacks chickens, ducks, and geese, and hence its damage to poultry farming is practically nil. It causes more damage to game by destroying valuable pheasants, hazel grouse, and fur-bearing animals (squirrels), but the damage is not great due to the relative scarcity of the cat. According to deer farmers, this cat sneaks into deer parks and destroys young sika up to one year of age (Menard, 1930). Referring to information obtained from former owners of deer nurseries, G.F. Bromlei (1956) reports that this cat attacks sika fawns up to one month of age only when the mother is not in the vicinity. A forest cat may be caught accidentally during a hunt for other animals. Usually it is shot with the aid of dogs, which tree it. It readily enters various traps baited with fresh hazel grouse. In the fur industry these pelts are included in the ‘‘wildcat’’ gro р, but’ identified as *‘Amur’’. Formerly, the pelts of this cat were exported to China where they were used to make collars, caps, and linings for ladies’ jackets. The range of the Amur cat in the Soviet Union is small and hence its total population quite low. The range is shrinking and the population in many parts dropping rapidly; the Amur cat could easily disappear altogether. Measures for its protection should be implemented, at least in those areas where the animal faces extinction. (A.S.) 270 356 JUNGLE CAT!” Felis (Felis) chaus Gildenstaedt, 1776 1776. Felis chaus. Gildenstaedt. Nov. Com. Acad. Petrop., vol. 20, р. 483. Terek. 1811. Felis catolynx. Pallas. Zoogr. Rosso-Asiat., vol. 1, p. 23, Terek. 1898. Felis chaus typica. De Winton. Ann. Mag. Nat. Hist., ser. 7, vol. 2, р. 291. For F. ch. chaus Giild. 1898. Felis chaus furax. De Winton. Ibid., p. 293, Jericho, Palestine. 1939. Felis chaus prateri. Pocock. Fauna Brit. India, Mamm., vol. 1, р. 298. Jacobabad, Sind, northwestern India.!® 1969. Felis (Felis) chaus oxiana. Heptner. Zoolog. Zhurn., 48, 8, 1259. ‘“Tigrovaya Balka’’ preserve, lower courses of Vakhsk [River], right tributary of Amu-Darya. (V.H.) Diagnosis Color of trunk and head monochromatic, without distinct spots or stripes; dark markings occur only on the tail and legs. Skull long, but palate not extended posteriorly in the interpterygoid vacuity (see Fig. 28); interpterygoid vacuity broad. Second premolar (first in row) usually present; antero-inner: cusp of carnassial tooth well developed and with a high pointed cusp. (V.H.) Description Size small, corresponding roughly to that of Caucasian forest cat [Felis silvaticus caucasicus], but somewhat larger in general. Some animals may be very large. In general appearance jungle cat differs somewhat from forest or steppe [wild] cats in that the former stands slightly taller on its legs and has a relatively short tail (Fig. 166). Tail length is less than half, usually about one-third of body length, and half the length of the hind limb (rarely equal '7Also called ‘‘chaus’* and sometimes *‘jungle lynx.’’** The latter name is artificial, simply a translation of the German name, and extremely unsatisfactory since this species is not related to lynx. The small tufts on the ears confused earlier investigators. Moreover, in old illustrations artists not acquainted with the live animal gave it a lynx-like appearance — extremely long legs and lynx-like ears. *Also reed cat and reed lynx —General Editor. '8shawiana and maimanah are sometimes included among synonyms for species in Russian fauna. These names should not be applied to the jungle cat (see p. 441 footnote and synonyms for cats of /ibyca group). ‘лолешом ‘КГУ Аа YOJOYS “PIND snvyo (51241) $25 ‘во э18ип[ ‘99т “Sty 358 to it). These proportions do not, however, reflect a similarity between jungle cat and lynx, as often stated. Head fairly broad in region of zygomatic arch and hence appears more rounded (spherical) than that of Amur cat. Ears quite long, relatively broad at the base, pointed toward the end, and set quite high. Small tuft of long hairs occurs on ear tips in winter. These hairs form an indistinct tassel ranging from 7 to 10 to 20 mm in length. Ears invariably project out of coat. Hair on cheeks not long (side whiskers not developed). Eyes fairly large and pupil vertical. Head or ‘‘face’’ of jungle cat altogether typical of small cats, and, like that of the forest [wild] cat, has nothing whatsoever in common with the lynx. Pelage throughout the body uniform in length, but in summer, hair on back long and forms a narrow and sparse crest of bristly guard hair all along the spine. The hair in this crest may be up to 50 mm long. In some animals this crest is weakly developed, however. In winter coat a crest per se is not seen, but the hair along the middle of the back is perceptibly longer. It is quite possible that the summer crest is formed of unmolted winter hair. The guard hair of the winter coat is relatively long and resilient and hence, in spite of the soft underfur, the coat of a jungle cat, compared to that of other small cats, is sparser and rougher. Neither in winter is it long and rich, but sparser, short and rough, as in summer. In winter about 4,000 hairs occur per cm? on the back and 1,700 on the abdomen. Extreme growth of top hair on the abdomen is a characteristic feature. The ratio of underfur to top hair on the back is approximately 12: 1 and only 4: 1 or 5: 1 on the abdomen. The top hair is very long; that of category I is 1.8 times longer than the fur. The average length of guard and top hairs of categories I to IV on the back is 61, 56, 47, 43, and 39 mm, and their thickness 89, 74, 52, 40, and 30 microns!°; on the abdomen the corresponding values are 50, 37, 30, and 26 mm (top hair of category IV absent), and 48, 30, 28, and 27 microns. The underfur is 30 mm long and 20 microns thick on the back and 20 mm and 21 microns on the abdomen (B.F. Tserevitinov). On the whole the body color of a jungle cat is monochromatic and devoid of spots. Dark markings and spots occur only on the legs and tail and sometimes on the abdomen. The general color of the winter coat (average type of coloration of Caucasian animals) is grayish-ocherous with an indistinct, very slight ripple consisting of light and dark top hairs. On average, more white hairs occur in this ‘‘ripple’’ and hence the coat has a grayish cast. Behind the scapular region a band extends along the back, which is initially narrow and indistinct but later becomes quite broad and turns a Measurements along the middle of the back, i., in the region of the *‘crest’’, are described. 359 deeper reddish-ocher color. Dark-colored hair is finely sprinkled in this band near the scapular region, but its density increases toward the rump. On the sacrum, and sometimes at other places, the density of dark-colored hair may actually form narrow fringes along the ocherous band. Flanks lighter in color than the back and admixture of top hair (small ripple marks) reduced; down the flanks, color merges rather abruptly into that of abdomen. Chest and stomach a vivid light ocherous shade with sections of white on chest and axilla; posterior third of abdomen and groin regions white. Chin, region of lower jaw, and throat also white. Basal part of neck and anterior part of chest a vivid light ocherous color. This field expands in the form of a cape into the region between the forelimbs. Outer side of hind legs identical in color to the flanks; outer side of forelimbs dull ocher; admixture of dark top hair in latter less than on hind limbs, or absent, and hence forelimbs appear brighter than hind ones. Two dark brown transverse stripes run into upper portion of inner side of forelimbs on an extremely light ocherous-white background, or a few spots to five or six short stripes of the same color may occur. Top of neck the same color as anterior part of back; sides of neck slightly lighter and approach color of the flanks. Uppermost part of tail the same overall color as the back or slightly grayer— ocherous stripe reaching base of tail does not extend onto Fig. 167. Young jungle cat. Trans-Caucasus. Photograph by A.P. Zhandarmov. ZS 360 it. Proximal portion of tail exhibits a strong admixture of black top hair; tip black with two or three black rings above it; first and second rings sometimes connected by a longitudinal band. On the head, between the ears and the occiput, color the same as on the back but more saturated and brownish. Upper lip and region of vibrissae bright white, or slightly ocherous, and stand out (characteristic features of the species) (Fig. 168). A dark brown spot occurs in front of the eye toward the nose, and a rather vivid ocherous field under the ear, behind it, and on the cheeks. Rest of the head same color as the flanks. Back of ears a vivid ocherous-red, but slightly brownish at the base; tip of ears brown or black, and tuft black. Inner side of ears covered with white hairs, the longest of which are pure white. Vibrissae white. Individual variability of winter coat relatively minor; coat either more grayish or more ocherous in general shade, and shape and color of band on back varies. Latter nay be short or long, very broad and indistinct, or Fig. 168. Jungle cat, Felis (Felis) chaus Gild. Bright coloration of upper lip a characteristic feature. Photograph by M.I. Obukhov. 274 361 relatively narrow, sharp, and rather vivid. Intensity of color of the underside of body also varies, but bright spot on neck and chest distinct. Brightness of color and outlines of spots vary slightly. Large number of spots (bands on forelegs invariably two), and poorly developed, indistinct ocherous spots may be visible at the boundary between color fields of flanks and abdomen, as well as on the abdomen. Color of summer coat far more variable than that of winter coat. On the whole it is far brighter and only rarely as dark and dense as the winter coat. General color tone of back varies from ocherous-gray to almost pure gray, with a slight dark- and light-colored ripple—gray tone may be quite dark or very light. Stripe on back, as in winter coat, varies in length and width and ranges from ocherous-brown to bright ocherous-rust. Color of underside of body less vivid and far lighter—white fields larger and ocherous tone paler. Sometimes, however, entire underside of body, except for chin, a uniform bright ocherous color, even exhibiting an orange hue. Spottedness somewhat more intense in summer than in winter; spots on legs and at border of color of flanks and underside more vivid, while spots on abdomen better developed and sometimes fuse into transverse stripes. Very light-colored and pale animals” obviously represent an extreme and very rare type of color variation in the summer coat. Top of neck light and grayish; only a relatively narrow gray band (almost equal in width to ocherous belt) extends down the back; sides, however, light ocher. Color may also be ocherous-gray with a rusty band. A narrow black stripe occurs on the tail which, as such, is white. Lower portion of tail very light-colored. Chin, throat, and chest between legs white, and area between chest and throat light ocher. Posterior part of chest and anterior part of abdomen pale ocher, and posterior part of abdomen and groin white. Spottedness noticeably rich. They are well expressed along legs and abdomen, or legs, flanks, whole of abdomen, and whole of chest covered with rust-brown (hind legs and flanks), brown (forelegs), and bright rusty (flanks and underside) indistinctly defined spots. Coloration in this type of spotted pattern is quite dramatic. General color of first coat in young animals (June, Kushka) dirty gray. A broad, poorly delineated dark brown-gray field extends down the back. Along the sides of this field, on the back, flanks, and abdomen, indistinct gray spots tend to form vertical rows. The field between them on the back and flanks is a slightly lighter gray. Underside of body light, pale yellowish- white, with yellow hue deepening in anterior part of abdomen and on the chest. Forelegs above the ulna rust-colored but dull; small blackish-brown spots occur below the ulna on both the outer and inner sides. Hind legs bear 20Мо. $ 14194; Aleksandriisk station, lower Terek; July 5, 1926; Yu.G. Heptner. No. $ 52701; Gasan-kuli at mouth of Atrek, Turkmenia, 1942; A.V. Samorodoy, KZMMU. See later section, *‘Systematic Position”* 362 brown spots and both upper and lower parts of feet rusty. Dorsal side of tail dark brown and underside light ocherous. Top of head same color, as the back and sides slightly paler. Field under the eye and to the rear of it, under the ear, ocherous-gray; upper and lower lips, chin, and throat white (see Fig. 167). Back of ear rust-colored, tip black, and ear tuft black and about 10 mm long. Juveniles assume an adult coat in their first winter. This species displays distinct geographic variation in coloration and is a good example of Gloger’s rule (Weigel, 1961). In addition to a general intensity of color in some subspecies, range and type of individual variation also seem to differ—gray and reddish-brown phases, together with transitional colors evident in the torso (F. с. affinis), darkening of color of the legs, and so on. Within the Soviet Union color variation is insignificant and mainly reflected in general hue and sometimes individual patterning. The skull of the jungle cat (Fig. 169) occupies an intermediate position between that of the peripheral form (in the Soviet fauna) of the genus —the Amur cat—and more specialized forms of the subgenus Felis $. str.— group silvestris, species silvestris. This similarity is exhibited in the slightly longish form of the skull in general and the brain case in particular, the comparatively narrow zygomatic arches, the form of the nasal processes of the premaxilla, and in other details. At the same time, the brain case in jungle cat is more expanded, roomier, and less extended posteriorly, the frontal area large, bulging, and with a small depression in the center, and the entire skull per se more convex (upper line of profile even above the arch is acute). Furthermore the orbits are relatively and absolutely larger; though set only slightly more forward, they extend slightly more posteriorly and take on an oval shape. The palate does not extend posteriorly to the same depth into the interpterygoid vacuity as it does in the Amur cat; in the jungle cat the posterior boundary lies at the level of the posterior margin of the molars (see Fig. 28). The interpterygoid space is broad, somewhat broader in front than behind, and the width of the choanal opening much greater than its height. The anterior width of the interpterygoid vacuity exceeds the distance between the inner margins of the auditory bullae. The teeth, especially the carnassial and third premolar, are very strong and massive. The anterior inner cusp of the carnassial tooth is well developed, with a fairly large pointed peak. The second premolar is usually present. Most of these features are developed to such an extent that in differentiat- ing the jungle cat from the Amur cat, the former comes somewhat closer to forest and steppe [wild] cats, and even to the caracal (see ‘‘Systematic Position’). Age-related changes in the skull are evident in the relatively larger volume of the cranium, lesser expansion of the skull per se and its more spherical shape, lesser development of the orbital processes, and absence 275 276 363 or poor development of the sagittal and lambdoidal crests. The proportions and structure of the skull of an adult animal are not seen in a juvenile prior to the middle of its second year (which means its second winter). The skull of females are usually lighter and somewhat smaller than that of males. Geographic variation in the skull is almost nil and seen only in insignificant differences in overall size. In body size, jungle cats in various parts of the Russian range do not vary much, and the data given for the region in the upper reaches of Amu- Darya (Termez and above, Table 7) are also applicable to the Caucasian population. Significant variations in weight (twofold or more in males and almost threefold in females) are attributable not only to significant individual and age-related variability, but also to seasonal changes and availability of food in different years. In the Amu-Darya delta the weight of males (13) in March-April averaged 7.9 kg and in October to December (4) 8.2 kg; the corresponding values for females (4) were 5.5 and (3) 6.5 kg (Reimov and Nuratdinov, 1970). Under favorable conditions these cats can become ex- tremely obese. Absolute mean weight of heart in males (21) 39.2 + 3.9 with a coefficient of variation of 34.2 and in females 35.8 + 4.7 and 29.3 respectively. Relative weight averages 4.7 + 0.25%o with a coefficient of variation of 17.0 and 6.1 + 1.0 and 36.7 respectively. Absolute average length of intestine in males (21) 183.9+5.9 (coefficient of variation 10.98) and relative length (for average body length of 76.5) 1 : 2.40; absolute length in females (15) 176.5 + 12.0 (coefficient of variation 10.82) and relative length (for average body length 68.8 + 2.2) 1 : 2.56. The absolute and relative weights of the heart, depending on degree of obesity, also reveal quite significant individual variations, often in different seasons of the year—males averaged from 39.7 to 34.6 g and 5.02 and 4.22%o, and females 40.4 to 29.2 g and 7.34 and 4.46%o (according to Reimov and Nuratdinov, 1970 with modifications; data for Amu-Darya delta). Skull measurements, based on data for the whole of the Russian range of the species, except the Amu-Darya delta, are given in Table 8. Data for different sections of the range are presented under “‘Geographic Variation’’. As in the case of body size and weight, the skull measurements of males significantly exceed those of females by an average of 15 to 16% with respect to the first category of characteristics, and roughly by 10% with respect to the second category. (V.H.) Systematic Position As briefly mentioned above, the jungle cat in skull structure (see Fig. 169) 364 ..ЧОЦЕЧЕД 91418090, 1029$ des ‘взер 1ецоштрре юд ‘лолеАв[\у ‘ИГУ Aq paysiwing [еиззеш рие (8661) лэузАшэцо "Гл JO вер UO paseg :эюм $506 0052 8195 РТ €L0°8 000‘ZT 000'S IT 14812 AA OL 08 09 OL SL 58 IL OT $1еэ JO 148 ЭН IST Z91 ОРТ ZI €91 OST OST и JOO} puly Jo Ч18иэ] 695 865 055 ZI 595 OTE 005 ZI ysugy пет, 859 058 595 ЕТ €9L 076 059 ЕТ 413иэ1 Apog W хеш ии И : W хещ иги И $э[еи1э1 ар\у 5этеш ару 121495тр [эло4е pue 7э1э1] eAreq-nwy лэ44п ayy woay ‘vuvixo snvys (J) “yf ed aysunfl Jo (8) 3481эм рие (WW) э715 Apog */ эЕТ, 9[< 365 `914е1 $141 UT papnyour иээа jou эльц (, цоцецел э{4е130э0,, и0Ц2э$ 99s) е|эр еАзе-пш\/ шо зреиние 10 ве :эюм ‘eAreq-nury 12441 wo Ато тенэе И ‘snseoned ay} шоц Ашо [eae 8'65 [РЕ 597 6 856 06 8'65 9 ZPUIWIRIOJ |еиалоелуиг иээм1э4 YIPIAA 9`ЕТ 9'РТ STI 8 6'РТ 6'SI СУТ 9 гоу JerssvuseD Jaddn jo чаи (эитиво YIM) 99E 16€ [РЕ 8 SOP OC 0'6£ IT MoI yoo} Jaddn до yysuaT OSE O'8E OEE L E9E TBE 0’ II YIPIM [9319101504 9°61 SIZ O9r 91 6172 0'95 L6l ИТ ра теталюлэзи] SCL [708 0'59 91 O'€8 0'06 ECL LI WIPIM эцешозА7 r'ZOr 901 816 LI Cell 0'05т сот LI yysua] [езе4оАрио Tell ЕСТ 0'<от ИТ € 971 "СЕТ УИ a yysua] 1591210) W xeul uid И И xeul ии Ц $91897 пру 5эеш [арх snvy? (С) ° ‘ео э[8чиГ Jo ззиэшолазеэш [[NYS °g эет, 9/5 366 (у) и ni С В — Ss 275 Fig. 169. Skull of jungle cat, Felis (Felis) chaus Giild. No. $ 5376, male, Zoological Museum, Moscow University. Lenkoran, Vel’, February 10, 1925. Sketch by N.N. Kondakov. 367 occupies an intermediate position between F. euptilura and Е. silvestris — libyca, i.e., European forest and steppe cats. In this species (superspecies) Е. silvestris, the jungle cat is closer to the subspecies group silvestris than to libyca. Present-day taxonomists usually place it in subgenus (of genus) Felis s. str. (Pocock, 1939; Haltenorth, 1953; and others). In spite of prevailing views, in this subgenus the jungle cat is a less specialized form (slightly extended skull, cranium, position of orbits, and so on). This fact also determines its position outlined above, although the structure of the palatal, choanal, and interpterygoid regions differentiates the jungle cat from the Amur cat more than from European forest and steppe cats (species— superspecies silvestris). The indisputable taxonomic proximity of the jungle cat to the latter does not permit its segregation into a special subgenus (Chaus) as done until recently and sometimes even now (Petzsch, 1968). In view of the reasons stated above, there is no justification for bringing the jungle cat closer to lynx, although, as will be shown below (in section on caracal), in certain features of skull and level of its specialization from the Amur cat —manul lineage, the lynx (subgenus Lynx) can be separated from subgenus Felis at the morphological level of the jungle cat. In general body build, which reflects basic features of the lynx group, the jungle cat nevertheless represents a true cat of the steppe or Amur cat type (traditional opinions about this animal in this respect are biased). The color of the jungle cat is distinctive and differs markedly from that of other cats of the narrow subgenus as interpreted here. However, the spotted color of the young and individual variations in the spotted pattern of adults point to the close relationship between the jungle cat and the species of true cats. An intermediate color type, between the Amur and European forest cat, is thus not seen, although the similarity of the monochromatic color type of the Amur cat [see above] to the color of the jungle cat is quite evident. It is significant that in captivity (zoological parks), jungle cats produce fertile offspring with domestic cats (Gray, 1954); a more extensive analysis of hybrids has not been made. There is less definite information about the mating of wild Indian jungle cats with domestic cats (Allen, 1938). Nothing is known about this in the Soviet Union. The above-mentioned differences between jungle and Amur cats in fact represent a unique set of distinguishing features between Amur and forest cats. Hence, with the information available at present, there is no justifiable basis for separating Р. euptilura from cats of the subgenus Felis $. str. It is possible that these differences also may be found in other species usually placed close to F. euptilura (bengalensis, viverrina, rubiginosa, and planiceps—Prionailurus group). 278 The color and some features of the build of the jungle cat are distinctive and have no transitional types. The position of the jungle cat is also important in that it bears significant features of similarity with caracal in craniological features. Being a wholly typical member in the direct line of specialization of Soviet cats from Amur to manul, it represents that level in this chain of forms from which, so to speak, the lynx (subgenus Lynx) branched off (see characteristics of the genus and ‘‘Systematic Position’’ of caracal). (V.H.) Geographic Distribution Jungle cats are found in forest regions and areas close to water in the Near East, Middle, and Southern Asia, and northeastern Africa. Geographic Range in the Soviet Union The range in the Soviet Union (Fig. 170) represents the northern periphery of the range of the species. It covers the southernmost regions of the country and is divided into two parts —Caucasian and Middle Asian—which unite in the south outside the Soviet Union. Because of its preference for river valleys and other areas with abundant water, often with forests, the distri- bution of the jungle cat is extremely uneven and boundaries of its range, especially in the Soviet Union, are very complex. Extensive areas in which the animal is absent have been delimited. In the northwest jungle cats live in the Volga delta at places 65 km north of Astrakhan (Dosang) and among the Pri-Caspian reeds for a bit east of the delta (Dengizsk region, Guryev district). From the Volga delta the range extends southward to the mouth of the Kuma, and along its lower reaches, in the form of a narrow strip corresponding to the width of the reed beds along the Caspian coast. From the mouth of the Kuma the cat is distributed along coastal reeds southward to Chernyi market. It inhabits floodplains and stream channels in the area between the Terek branch running north to Chernyi market and branches [of the delta] running east into Agrakhansk Bay. Along the Terek it extends into the forests and thickets of the upper valley as far as Parabochev forest and Shelkozavod (Shelkovaya) station, and even to Mozdok and along the Sunzha to Groznyi. In the expanse between the Terck, in the foothills [of the Caucasus | and on the seacoast southward to Makhachkala (Dagestan plains) the cat is quite uniformly and extensively distributed, some distance from the sea, along the floodplains of the Terek, Sulak and its tributaries, and along lakes and shrubby thickets. It reaches as far as a line between Shelkovaya and Khasavyurt, or perhaps even farther westward. 279 278 369 To the south of Makhachkala the jungle cat is found along the Caspian coast to the Iranian border. Where reed thickets are absent this cat lives in shrubby thickets in the coastal belt or in foothills, rising up to heights of 300 to 400 m, and not more than 800 m, above sea level, but up to 1,000 m in the Trans-Caucasus. This cat is extensively distributed in the lowlands of the eastern Trans- Caucasus, along the Kur’ and Araks valleys and Kur’ tributaries, and among lake regions in the steppes. The range there is bounded by the foothills of the Great and Little Caucasus. In the Kur’ valley it extends almost to the Lake Balkhash Fig. 170. Reconstructed range of jungle cat, Felis (Felis) chaus Giild. in the Soviet Union (scale in km). Question marks pertain to the lower Chu and the Ferghana valley, information concerning which is dubious. V.G. Heptner. 370 Suram range (Gori) along the lower zone of the mountain and in the foothills of the Glavnyi [Caucasus] range and the Alazan basin, and runs in the west roughly to the meridian of Tbilisi. Along the river valleys of the Kur’ basin extending down from the Little Caucasus (Terter), it extends at places quite deep into the mountains. The cat lives in the Araks valley from its lower reaches up to Megra ravine (a spur of the Zangezur range on the Araks, slightly east of Ordubad) and, after several breaks (Southeast of Yerevan at the boundary of Nakhichevan Autonomous Soviet Socialist Republic), from Arazdayan to the Oktemberyansk region (west of Yerevan). It even penetrates there deep into the mountains along some tributaries (Akera and Okhchichai). References to the occurrence of the jungle cat in the Kuban (M. Bogdanov, 1873; Ognev, 1935; Formozov, 1946) are erroneous; only the forest [wild] cat (Е. silvestris?!) occurs there. Information about its occurrence at the mouth of the Ural [River] has not been confirmed and is evidently erroneous. In Middle Asia the jungle cat lives in the extreme southeastern comer of Turkmenia along the Atrek from its mouth (Gasan-kuli) and its tributaries, the Sumbar and Chandyr, and along the Tersakan (Khodzha-Kala). The animal does not penetrate deep into the mountains (along the Sumbar hardly beyond Kara-Kala). Throughout the rest of the Kopet-Dag this cat is evidently absent and also not seen in the adjacent plains. It does not go into the [desert] sands and references to its occurrence west of Uzboi (Yashkha and Toniatan; G.P. Dement’ev, 1955) are erroneous. Farther east the cat is found all along the Tedzhen, probably entering from Iran, and the entire Murgab. It is absent or rare along the Kushkha. In the Amu-Darya delta this cat is distributed to the [Aral] sea, and up the river to the mouth of Kyzyl Su and slightly beyond (to Chubek, south of Kulyab), 1.е., all along the river wherever reeds and shrubby thickets grow. Along the left bank of the river, beyond the delta, the cat is encountered only in its valley (desert closely approaches it). Only along the Kelifsk Uzboi and at the head of the Karakum canal does it deviate slightly from the Amu-Darya valley (probably the canal later leads the cat deep inside the desert). The jungle cat is extensively distributed along the right tributaries of the Amu-Darya; along the Kyzy] Su and its tributary the Yakhsu the boundary proceeds slightly above the Kulyab, runs along the Vakhsh a little above Kurgansk-Tyub (where the river flows out of the mountains), and along the upper Kafirnigan. It reaches the Gissar valley, in the east slightly beyond *1 4 specimen from Grivensk station (Zoological Museum, Moscow University), labeled jungle cat, belongs to this species. 280 ЗЕ Dushanbe (to Ordzhonikidzeabad), and along the Surkhan-Darya valley and the lower Tupalang. In the mountains between the above-named rivers it does not occur. Probably it lives along the Shirabad. It is distributed along the Zeravshan, where it is met with especially around Samarkand, and the boundary runs upstream along the river at least to Pendzhekent; it evidently lives on the Kashka-Darya. The animal occurs throughout the Syr-Darya, from the mouth of the Angren at the [Aral] sea, throughout the lower Chirchik and Angren, and the plains of the Tashkent region. Data are not available for the Ferghana valley. The cat apparently occurs along the lower reaches of Chu but these data require confirmation. References to the occurrence of the jungle cat in Ustyurt (Brandt, 1852; Bazhanov, 1951), not only in the central region, but also between the northern Chink and the Bol’shoi Barsuk sands, are erroneous. The cat lives around Lake Sudoch’ in the extreme northwestern part of the Amu-Darya delta, and short transgressions northward from there along the west shore of the Aral Sea should not be totally ruled out. The range of the jungle cat, in spite of the animal not being subjected to special intensive pursuit, and being known to survive well under certain conditions in cultivated areas (flood- and sprinkler-irrigated lands) undergoes considerable changes in certain places. This is explained by its characteri- zation as a stenotopic species. In recent years, as a result of the drying up of the Atrek and the disappearance of reeds along the river and associated water sources (Lake Delili), the entire reed bed fauna has vanished from this area, and with it the jungle cat. In southern Tadzhikistan the cat has disappeared at several places as a result of clearing of the tugais [bottomland forest and shrubland]. This has brought about changes in the distribution of the species along the Murgab. Contrarywise, as a result of irrigation and the cultivation of some regions in Tadzhikistan which until recently were deserts the jungle cat has reap- peared for example in the Vakhsh valley. There is reason to believe that the jungle cat, a normal inhabitant of the whole of the Syr-Darya valley, disappeared in the last decade from its lower course, or at least became rare, even though changes in the environment did not occur which would preclude its existence in this habitat.72 ?Вапое based on data of М. Bogdanov, 1873; Zarudnyi, 1890 and 1915; Dinnik, 1910-1914; Satunin, 1915; N. Smimov, 1922; S. Naumov, 1927; G. Nikol’skii, 1930; Ognev, 1935; Flerov, 1935; Shnitnikov, 1936; Gureev, 1937; Dobrokhotov, 1939; Heptner and Formozov, 1941; Vereshchagin, 1947; Kuznetsov, 1948 and 1948a; Chernyshev, 1948 and 1958; Bazhanov, 1951; Dal’, 1952 and 1954; Sludskii, 1953; Heptner, 1956; Nur-Gel ‘dyev, 1960; Shukurov, 1960; Ishunin, 1961; Alekperov, 1966; О.Р. Bogdanov, 1964; and others; and original data of A.A. Sludskii and V.G. Heptner (Turkmenia). 281 52 Geographic Range outside the Soviet Union The range outside the Soviet Union (Fig. 171) covers lower [elevations of] Afghanistan, Iran, Asia Minor (west and south), Iraq, parts of Syria, Palestine, Egypt (Nile delta and the lower course to Kep at 28° N. lat. and along the coast for 250 km west of Alexandria), and also Baluchistan, Kashmir, the whole of India and Sri Lanka, Nepal, Burma, Indochina (but not Malacca), and Yunnan.” Beyond the limits of its present range the jungle cat is known from the Holocene deposits in Germany and Switzerland (?), among relicts of the Azilian and Tardenoisian periods in the Crimea (Shan’koba) (V.I. and V.I. Ототоу, 1937, after Pidoplichko, 1951), and in Slavic settlements of the eighth to the thirteenth centuries, in Poltava territory (I. Gromov, 1948). That these remains belong to the jungle cat has justifiably been doubted (Vereshchagin, 1959). The identification of a species is hardly possible from such remains as were recovered. No other data are available about the occurrence of the jungle cat in the southern European part of the USSR. The jungle cat, at least in the Caucasus, is evidently a recent immigrant from southern Asia (V.H.). Geographic Variation Geographic variation in the jungle cat, in spite of its relatively small range, is quite considerable. Usually eight subspecies are recognized (Ellerman and Morrison-Scott, 1951 and 1966), excluding maimanah (see below; Pocock, 1951); or with the inclusion of the irregular, recently described form valbalala (Sri Lanka—9 (Weigel, 1961). Differences mainly pertain to color (its overall intensity, color of extremities, degree of variability, and so on), which conforms to Gloger’s rule (Weigel, 1961); also, extremely massive teeth have been described in one form (furax) (Pocock, 1939) but this feature is not fully convincing as judged from the material available with me. There obviously exist some differences in the overall size of some of the subspecies. For Soviet forms Bergman’s rule is also applicable. In spite of all this the *2By placing Е. shawiana among the forms of Е. chaus, Pocock (1939) included Chinese Turkestan (Yarkand) in the range of the jungle cat. In this respect he was followed by some western zoologists in particular Ellerman and Morrison-Scott (1951). This misplacement, as mentioned before (see synonyms of jungle and steppe [wild] cats), is evidently based on a misunderstanding of the data of collections, since jungle cats do not occur farther east than the Syr-Darya and perhaps the Chu. There is no positive information about the occurrence of this species in Yarkand; its occurrence there is highly improbable. Jungle cats have reportedly been sighted in northwestern Africa (Heim de Balzac, 1936); this information is not confirmed. 280 373 аи SV ATES | \ 3 ЕС 1000 0 1000 2000 3000 4000Кт ря SRR CSSD сити SEE | Fig. 171. Species range of jungle cat, Felis (Felis) chaus СШЧ. (scale in km). Question mark refers to lower Chu. V.G. Heptner. р distinguishing features of some subspecies have not been clearly defined and hence the actual number of geographic forms will probably prove fewer in future studies. Two subspecies are known in the Soviet Union. 1. Caucasian jungle cat, Е. (F.) с. chaus Giildenstaedt, 1776 (syn. catolynx, typica). Color relatively dark and traces of a spotted pattern relatively distinct (‘‘Description’’ given above pertains to this form). Large-sized form. Adequate data on body size and weight are not available but, judging from skull size (Table 9), they are not inferior to the form oxiana (Table 8). The weight of three adult males was 6,200 (June), 7,400, and 5,700 g; two females weighed 5,560 and 6,800 g (Azerbaidzhan; 374 ‘papnjour (ешошузат,) 3eq-iadoy ay) шо э[ешэу эцо рие эеш эио ‘(ст6т) UlUMIeS УМ Woy вер рие “(ssoUaIDg Jo Ашореэ\у “шпэзпи( [е1Зо[ю07, цезеопео ay) и! [енэзеш ‘се6Т) ^эиЗо 1$ JO вер ‘Аизлэлш Г MOOSOY] ‘шпэ5пу [291801007 aU} Ul [е1э3е| :э30 СД orl SCL 8 бт 6ST СУТ 9 4100} [е1ззеилео Joddn jo ц18чэ] 9`96 T'6€ LYE 8 S 0b 0’ O'6E т (ощиво чим) мол 4300} Jaddn до yy3ue7] OSE O'8E OEE L €9€ TBE OE п UIPIM 19170304 T'0Z LENG СЗТ L 917 0'95 SIZ Il уурим теалолэ и O7L SOL 789 L 568 0`06 0'6L п Чурим эвешо8А7, 850т YOIT 786 8 SPIT 0'02T TEOL от yysua] 1езеаоАриоЭ 6 СТ 8901 8 S971 OZET OT Il 433091 35212210) И хеш ии и И хеш и U Ssoxopuy so[ewoj ру 5эеш уару ЕЕ И S88 ЕС Е Е ee $$$ D000 5172 *2 ©) “ye a[sunf цебеопе о) эЦ} JO ззиэшоливаш [[NYS “6 242 L C87 282 375 А.А. Sludskii); an old female (Astrakhan’ preserve) weighed 13 kg (Dobrokhotov, 1939). This subspecies is found in the Volga delta, the Caucasus, and Kopet- Dag. No extension of the range eastward into Turkmenia is known. Animals from Kopet-Dag (KZMMU) in winter coat are somewhat lighter than Caucasian animals and present a somewhat transitional condition between this and the next subspecies; they should nonetheless still be considered the nominal form. Outside the Soviet Union this subspecies lives in Iran, Turkey adjacent to the Trans-Caucasus, Iraq (Mesopotamia), Syria and parts of Turkey adjoin- ing it, and Palestine (?). The nominal form is distinguished from almost all the other subspecies by its much larger size. In all the forms except furax and nilotica, the greatest length of skull from the published, though scanty, data (Pocock, 1939 and 1951) does. not exceed 117, 120, and 127 mm respectively, and the body length of one does not exceed 80 cm. However, for the form furax (Iraq, Syria, and Palestine) the greatest length of skull attains 131 and even 138 mm. Evidently it is identical with the nominal form; moreover most reports are from places quite close to the area where the nominal form is distributed (Baghdad; Pocock, 1961), and a large part of its range lies in the same natural region of the Near East as does the Caucasian form. The form from the lower Nile (иНойса— 131 mm) is also evidently close to the nominal form in size but will not be discussed here. 2. Turkestan jungle cat, F. (F.) c. oxiana Heptner, 1909. Color of trunk lighter than that of nominal form, with a lesser growth of black top hair, a more vivid band on the back, and more white hair on the underside. Underside of flanks very bright and ocherous, spots on legs fainter, and traces of spots in other parts of the body less distinct. Large-sized animals, evidently not smaller than the nominal form (see Tables 9, 10, and 11). The average body length of adult females from the Amu-Darya delta (15) is 668 + 22 mm and average weight (15) 6.08 + 0.13 kg (Reimov and Nuratdinov, 1970). The relationship of this form to that described from India is not clear. It is possible that it may be close to the form prateri, which occupies Sind and the region southeast of it. The two are hardly identical, how- ever. The type locality of this form (Dzhakobabad) is a straight-line distance of about 1,000 km from Termez on the Amu-Darya and beyond Hindu Kush. Furthermore the nominal subspecies is found in Iran and Baluchistan 376 [УЕ 81$ [08 т бот ЕССТ хеш S97 0’9т 0'59 S16 OCOL ии $этеэу Дару NN ON NaN = "L 219], Ul рэ}иэ$214 иээд элец UOISAI $11 шозу зрение JO 348тэм рие э71$ Apog :aj0N 8'СЕ УС cB TTL 9SCI W OVE СТС 788 Ltt 7 6CI хеш затеи зару сие о NOTING VUIWILIOJ [e}IqIovIJUI иээ/3э4 эоие1$1@ YIPIM Гезтатолэ и] рим эцешоЗА7 Ч3чэ1 |е5еа01АриоЭ 43343] 15232910) зэхэри] (SS6T ‘^эЧ$АилэЧо jo вер) еАле(-пигу Jeddn ‘numixo *2 СН) °Я Ged зип Г ay} Jo $цошола$вош [[NYS °0Т [42.1 <8С 283 ВИТ Table 11. Average body size (mm) and weight (kg), and skull measurements (mm) of male jungle cats, F. (F.) с. oxiana from Amu-Darya delta (from Reimov and Nuratdinov, 1970, with modifications) Indexes n Males Greatest skull length 17 123.8 = 2.06 Condylobasal length Я 112.9 = 1.84 Zygomatic width ily) 81.0+ 1.46 Interorbital width 17 20.9 = 0.46 Body length 21 755.0 = 32.00 Weight 21 8.9 = 0.56 Note: The average skull measurements of females published by these authors are not given since they are clearly exaggerated and not representative due to the small size of the sample (five animals) (V.H.). (Pocock, 1951). Moreover, the Sind form is notably smaller than the Tadzhik.”4 The following forms are usually recognized from some parts of the range outside the Soviet Union: 1) F. (F.) с. furax de Winton, 1898— Mesopotamia and Syria (see above under the nominal form); 2) F. (F.) c. nilotica de Winton, 1898—Egypt; 3) F. (F.) с. affinis Gray, 1830—Himalayas from Kashmir to Sikkim; 4) F. (F.) с. kutas Pearson, 1832—northern India from Bengal to Kutch; 5) F. (F.) с. prateri Pocock, 1939—Sind and the area to the southeast; 6) F. (F.) с. kelaarti Pocock, 1939—Sri Lanka and southern India south of Krishna River; 7) F. (F.) c. fulvidina Thomas, 1928—Indochina and Burma. (V.H.) Biology Population. The jungle cat is rare in the extreme north of its range, in the Volga delta. It is rare and sporadic on the northwest coast of the Caspian Sea. On the lower Terek and southward, to the boundary with Iran, it is *4The name maimanah Zukowsky, 1915, assigned on the basis of a visual examination (!) of the skin from Maimana in northern Afghanistan described by Scully in 1887, could probably be given to this form on the basis of marked similarity. Ellerman and Morrison- Scott (1951 and 1966) assign this name to the jungle cat with doubt (?). However, it refers to the steppe cat (see p. 442). 378 common, and at places abundant (lowland regions of Dagestan and Azer- baidzhan, and in parts of Georgia and Armenia). In Azerbaidzhan, until quite recently, some 3,500 of these cats were caught annually. The animal is now common but was formerly abundant at places in the southern half of Turkmenia (along the Atrek, Sumbar, Tedzhen, and Murgab). It is abundant on the Amu-Darya and its delta and on the southern coast of the Aral Sea. It is common and at places abundant in the tugais of the Pyandzh, Vakhsh, Kyzyl Su, Kafirnigan, and other tributaries of the upper Amu-Darya. It is particularly abundant on the Amu-Darya along the Afghan border. In Tadzhikistan up to 300 cats are caught annually. In Uzbekistan jungle cats are extremely common in the valley of the Amu-Darya and its tributaries (Surkhan-Darya, Kashka-Darya, and others) and in the Zeravshan valley. In recent years it has been rare in the Syr- Darya, Chirchik, and Angrensk valleys. In Syr-Darya and Chu in adjacent Kazakhstan this species is no longer met with, although earlier authors considered it common there. Habitat.» The jungle cat is a typical inhabitant of reed thickets, im- penetrable thorny bushes, dense, wet lowland forests around lakes, river floodplains, and along sea coasts. In open steppes or deserts it is encountered rarely and never ascends high into the mountains. The above-mentioned regions abound in swamp and water birds, francolins [Francolinus], water or complex-toothed rats [Nesokia indica], mouse-like rodents, several species of reptiles and amphibians, and also fish. The cat is assured of abundant prey in these hunting grounds. In the Volga delta it lives in reed and cattail thickets, which alternate with i/mens [Volga delta lakes], and are dissected by abundant small rivulets—eriki. Willow shrubs grow on the banks of these shallow channels. While hunting, the cat sometimes reaches the coast. It lives in reed and cattail thickets with a large number of breaks in the lower Kuma, Terek, and other rivers of the Caucasus and Trans-Caucasus and also in the large system of lakes (Sary Su and others). In Armenia the animal lives, for example, in the Araks valley in and adjacent to reed shrubs at a height of 800 to 970 m above sea level (Dal’, 1954). On many of these water sources, reeds and cattails grow in the water in the form of a strip 1.0 to 3.0 km wide. These bushes surround the pools like a dense wall, isolating them from the adjoining steppes. There are often bogs and floating islands in these lakes. The cat lives in such thickets year-round and ventures into adjoining sections only occasionally and briefly, for example the steppe. Hither it sometimes emerges in winter to hunt rodents and steppe game. This cat when hunting enters the water and Photographs given before of habitats of Turanian tiger, except for Semirech’e (Ш River) also pertain to this species. This cat and tiger are sympatric almost everywhere. 579 284 Fig. 172. Habitat of jungle cat in Astrakhan preserve. Volga delta. June, 1950. Photograph by A.A. Sludskii. is an excellent swimmer. In the Trans-Caucasus it is often found in breaks in the reeds and bogs 1.0 to 1.5 km from the bank. It swims through large pools not only in summer but often even in winter. It is so accustomed to water that it often rests in breaks among reeds far away from the bank and sometimes even sets up its den there. When chased by man or dog, it usually jumps into the water and escapes by swimming. In addition to reed thickets jungle cats often live in almost impenetrable growths of blackberry and clematis growing near water, and in tugai forests of turanga [Euphrates poplar] and blackberry. In tugais it is quite common, for example on the Kur’. In the Lenkoran lowland and also in other places it is not infrequent in the shrub zone of the foothills and in reeds by the sea. On the coast it lives in winter wholly on emaciated ducks and coots and also on injured animals. Tracks of this cat are encountered everywhere along the outer edge of the thickets and on open sandbars. Here the remains of its prey are often encountered, i.e., birds torn to bits. (Yu.A. Isakov). In the Lenkoran lowland, however, cats are abundant in gardens, where they live in hedgerows consisting of white willow densely intertwined with climbing, thorny vegetation. At places these thickets are so dense and exten- sive that part of a wild garden may be transformed into an impregnable fort 380 286 Fig. 173. Alder swamp in Agri-chai valley. Biotope of jungle cat. Azerbaidzhan. December, 1957. Photograph by N.N. Rukovskii. (E.P. Spangenberg). When living close to man, the cat often intrudes into villages and steals domestic fowl. This cat has been found several times in barns, around dwellings, and other places. Jungle cats have sometimes been seen in the Trans-Caucasus in places that are pure steppe, several kilometers away from water sources (Smirnov, 1922). In Middle Asia this cat also inhabits reed thickets and tugais. How- ever, in Tadzhikistan in March and April, when desert hares [Lepus tolai] and pheasants migrate from the tugais into the adjoining desert to fatten up, the predator follows them there (Chernyshev, 1958). In the Amu-Darya delta it lives in extensive reed thickets, often flooded in summer, and is frequently sighted in bogs and breaks among the reeds a few kilometers from the banks, or on the shore (A.A. Sludskii, V.S. Pokrovskii). In Middle Asia this animal occurs in newly irrigated sections of the desert and inhabits thickets along the banks of canals and major irrigation ditches, which are reminiscent of miniature tugais. Pheasants, gerbils, hares, complex-toothed rats, and domestic mice share these habitats with such predators as jackal, steppe cat, and jungle cat. On the other hand, where the tugais have vanished the jungle cat has Stee ERAT ed ее ее ие те пе еее ое meeuceeategegnmmene eter ear re 286 Fig. 174. Unfrozen opening оп a river where the jungle cat hunts for wintering ducks and picks up injured ones. Severe, abundantly snowy winter, 1949/1950, in Trans-Caucasus. Kyurdamir, Azerbaidzhan. Photograph by S.S. Turov. become rare. For example, at places in Tadzhikistan where cultivated land has largely replaced tugais, this cat has become a rare animal. In the Gissar valley it survived only in infrequently occurring ‘‘islets’’ of reed thickets in the area of Gissar (Chernyshev, 1958). As a result of the destruction of its biotopes in recent years, the distribution and population of this cat in Azerbaidzhan has decreased rapidly (Aliev and Nasibov, 19066).?° As an inhabitant of lowlands, the jungle cat ascends only up to 600-970 m above sea level in the Caucasus and Trans-Caucasus (Dinnik, 1914; Vereshchagin, 1947; Dal’, 1954), to 900-1,000 т at Dushanbe in Tadzhikis- tan (Chernyshev, 1948), 1,800 m in Iran, and even 2,400 m above sea level in the Himalayas (Blanford, 1888-1891). Specialized for predation on animals closely associated with unfrozen water sources or those frozen only for a brief duration and not annually, the chaus finds optimum living conditions only in the far south of the Soviet 26Тре reports of С. Radde, A. Walter, [and W. Blasius]* (1889) that in the southern regions of Turkmenia, altogether devoid of shrubbery thickets, jungle cats were present and inhabited crevices and small caves in gorges on the banks of upper Murgab requires confirmation. These investigators report finds of jungle cat in such an environment on April 8 to 20, 1887 at Takhta-Bazar. *Omitted in Russian original —Sci. Ed. 287 382 Union. The northern limit of the massive wintering of waterfowl in the Soviet Union roughly coincides with the January isotherm of 2°. Evidently this isotherm also serves as the northern boundary of optimum conditions for the cat. It is not well adapted to subzero [Celsius] temperatures and snow cover. Of all our wild cats, the chaus has the shortest, coarsest, and sparsest hair coat, which poorly protects the animal from cold. Hence, in the northern part of the range during unusually severe and abundantly snowy winters this cat perishes from cold and hunger. In the severe winter of 1941/1942 in Astrakhan preserve one cat came to a stable on February 1 and three dead ones were found in the Terkhizbenka River. The animals were emaciated and their intestines empty (Yu.A. Isakov). The absence or extreme rarity of this animal in Kazakhstan and the southeastern European sector, in spite of extensive reed thickets and swamps, is explained by the above ecological features. In addition,.snow hinders cats venturing high into the mountains. Food. On the western shore of the Caspian Sea this cat feeds on water voles [Arricala terrestris], small mouse-like rodents, European hare [Lepus europaeus |, ground squirrels [Spermophilus], wild boar (small sucklings), insectivores (shrews), and in recent years the acclimatized nutria. It attacks ducks of various species, coots, moorhens, pheasants, francolins, gray part- ridges, and smaller birds; it also destroys fledglings and eggs. It eats turtles, snakes, lizards, and fish (mainly sazan). When living close to villages, it often steals chickens, ducks, and geese. It eats fresh carrion. Its main prey year-round are rodents and waterfowl (ducks and coots), especially in winter, followed by pheasants, francolins, and fish. In several regions the cat lives almost exclusively on waterfowl. How diverse its prey is can be judged from the following examples. In the stomach of a chaus caught on the lower Terek (June) the remains of ten ground squirrels were found, and in the stomach of another caught in winter in Dagestan were some shrews swallowed whole - (Heptner and Formozov, 1941). The stomach of a large male from Kyurdamir (Azerbaidzhan; June) contained two water voles, the scales of a sazan, and about 800 g of the meat of a small wild boar; the stomach contents of a cat from Kyzyl-Agach preserve (August) showed one water snake [Natrix tessellata or sipedon|], a pochard [Athaya ferina], and three house mice; the stomach of another caught in December contained a pochard; and that of acat from Karayaz (Georgia; February) had one blackbird and two common voles (Vereshchagin, 1942). A marsh turtle was also found in the stomach of one cat. Jungle cats often hunt for nutria; for example in Karayaz (Georgia) they and jackals act as the chief agents limiting the nutria population. Even from farms fenced with a wire net to a height of 2.5 m, these cats stole 147 young and adult nutrias during the three winter months of 1947 (Vereshchagin, 1950). A particularly large number of nutrias were killed by cats in the 288 383 Fig. 175. Reed floodplains in ‘‘Tigrovaya ВаКа`” preserve. Habitat of jungle cat. Tiger, wild boar, Bukhara deer, and pheasants are also found here. Lower Vakhsh, Tadzhikistan. October, 1960. Photograph by A.A. Sludskii. unusually severe winters of 1948/1949 and 1949/1950. In these winters, because water sources froze over, the cats caught nutrias hiding in thickets or running away over ice. The predator evidently catches mainly young nutrias and rarely attacks adults. A cat was once observed attacking a large nutria, which made no attempt to escape from its enemy, nor to dive into the water, but protected itself, and sometimes itself attacked. After 10 to 15 minutes, the predator was compelled to withdraw, and after meowing swam away into the reed thickets (Pavlov, 1953). At such times [severe winters] cats kill many waterfowl weakened by hunger, the more so because they are concentrated in large numbers in a few frozen* stretches (Е.Р. Spangenberg). In the region of Kyzyl-Agach Bay in the severe and abundantly snowy winter of 1956/1957, little bustards [ Tetrax tetrax| and waterfowl wintering there had become greatly enfeebled and thus fell easy prey not only to foxes and jackals, but also to jungle cats (T.A. Adol’f). *Unfrozen? — Sci. Ed. 288 Fig. 176. Habitat of the jungle cat in ‘“Tigrovaya Balka’’ preserve. Lower Vakhsh, Tadzhikistan. April, 1966. Photograph by G.N. Sapozhnikov. While hunting for waterfowl, water voles, and nutrias, cats will often swim from bog to bog, sometimes across long stretches, not only in summer but even in winter. In hunting, upon reaching an opening among reeds, the cat simply lies in wait for any prey; noticing one nearby, frequently jumping directly on a swimming duck or nutria in the water (Kashkarov, 1932; M.P. Pavlov). In January, 1949, along a backwater of Lake Tomkin (Tadzhikistan), a chaus jumped from the bank onto ducks swimming in an opening in the ice and caught one of them (Chernyshev, 1958). In the summer of 1951, near Kyurdamir (Azerbaidzhan), as a result of intense drought many lakes dried up considerably. The water receded from the shore almost a kilometer, exposing extensive mud-flats; holes with water and fish in them remained here and there. In the drying !ake from dawn to 7:00 to 8:00 a.m. cats hunted regularly, catching fish in these pits. Usually the predator covered the mud-flat between the bank and the hole in long leaps, and on seizing a fish, and quickly ran back into the reeds. The cat pounced in a long leap on sazans weighing 200 to 500 g moving in the shallow water, snatched one of the fish, and retreated rapidly to cover. Cats also preyed on fish at the end of the day, often even until sunset. In 385 this same area and year they also stole water voles from traps (Yu.A. Gerasimov). In regions where autumn and winter hunting for waterfowl is well organized, wounded birds and those weakened during overwintering often 289 constitute the main quarry of the cat. In southwestern Turkmenia the main 290 Table 12. Results of analysis of feces and stomach contents of jungle cats in Middle Asia (percentage of total number of observations) Region, season, and material Right tributaries Floodplains of Lower of upper upper Amu- Amu-Darya? Amu-Darya Darya, Aral- (Tadzhikistan)! Paigambar Island? Food — Summer Winter — 100 samples 33 samples (including 33 419 53 (including six stomachs) samples samples — stomachs) Mammals 57.2 —- - 67.1 Shrew 2.2 0.6 — 4.1 Desert hare [Lepus tolai 15.2 100 10 13 tibetanus] Rodents — 850 920 63.0 Small jerboa [Allactaga — — — 1.6 elater| House mouse [Mus musculus] го 4.0 13.0 232 Complex-toothed-rat [Nesokia] 4.4 110 44.0 1.3 Midday gerbil [М. meridianus] — 75.0 26.0 Red-tailed gerbil [М. erythrourus]| 4.4 — — Crested gerbil [М. tamariscinus] — — — IWS) Great gerbil [Rhombomys]| — — — 4.1 Mole vole [E//obius| = — — 2.6 Trans-Caspian vole — — 123 [М. transcaspicus| Muskrat [Ondatra] — — — 8.2 Мила [Myocastor] — — — 1.3 Rodents, not further identified SED — — 4.1 Boar (suckling) — 0.4 — — Jackal [Canis aureus] 22 — — came Birds 36.2 120 44.0 3165 Pheasant [Phasianus] 18.7 3.0 17.0 Duck 2.2 — — 2.6 Pochard [Athaya ferina] DD — — = Coot [Fulica ата] ial wa nite Needy 386 Table 12 (Contd.) Food Cormorant [Phalacrocorax] Black-necked sheldrake [Red-headed pochard, Netta rufina? | Harrier [Circus] Birds, not further identified Pheasant eggs Reptiles and amphibians: Lake frog [Rana ridibunda] Steppe agama [A. sanguinolenta] Box lizard [Eremias] Bluntnosed viper [V. labetina] Desert tortoise [Тезшао horsfieldi| Eggs of turtle Fish: Sazan Gambusia Arthropoda: Scorpions Phalangids Insects Plant food: Oleaster nuts Green stalks of reeds Green stalks of sugar cane !V J. Chernyshev (1958). °G J. Ishunin (1965). 3A.M. Allayarov (1964). Region, season, and material Right tributaries Floodplains of Lower of upper upper Amu- Amu-Darya* Amu-Darya Darya, Aral- (Tadzhikistan)! Paigambar Island? — Summer Winter — 100 samples 33 samples (including 33 419 53 (including six stomachs) samples samples stomachs) Ahi — — — — — — 1.3) — — — 2.6 of} 12.0 44.0 29.2 — 3.0 — — 4.4 16.0 — 1.3 1.1 — — — — — — ES Uhl 8.0 — — 22 7.0 — — — 1.0 — — — 0.4 — — 35 — — — 2.2 — — — 3.3 — — — 33 — — — — 3.0 — — — 1.0 — — — 23.0 22.0 — 14.3 3.0 30.0 — 55 1.0 17.0 — 4.4 — — — 3.3 — — — 387 289 Fig. 177. Тиса! along a dried up tributary. Habitat of jungle cat in ““Tigrovaya Balka’’ preserve on Lower Vakhsh. Jackai, tiger, and pheasant also live here. October, 1960. Photograph by A.A. Sludskii. prey are rodents and birds, especially overwintering waterfowl, which are caught during the night, and francolins. я. food of the jungle cat in Middle Asia, see Table 12: It can be seen from Table 12 that in Tadzhikistan and Uzbekistan jungle cats feed mainly on desert hares [Lepus tolai tibetanus], rodents (63-92% of samples), and birds (12-44%). Jackal remains in feces are evidently explained by the consumption of carrion. There is a definite place for plants in the diet of cats, especially oleaster fruits (Eleagnus angustifolia), the nuts of which are regularly encountered in feces. The food regime varies very little in different seasons but hares, complex-toothed rats, and pheasants predominate in the summer diet. In the Amu-Darya delta, in addition to birds and complex-toothed rats, the cat hunts acclimatized muskrats by watching at its dwelling or digging up the house; it destroys the constructions of this rodent more vigorously than does the jackal. In the muskrat harvest season cats approach very near to trapping parties and pick up discarded muskrat carcasses (A.A. Siudskii and V.S. Pokrovskii). 388 In Afghanistan the stomachs of three chaus contained the remains of 19 house mice, 3 complex-toothed rats, and 9 lake (?) frogs (Neithammer, 1966). The stomachs of cats have contained up to 1,200 g of food. In zoological gardens they are fed 720 g of meat with bones daily in summer, and 870 g in autumn and winter (Obukhova and Shakhnazarov, 1949). Home range. The size of individual home ranges are not known. Judging from tracks, they cover in one night no more than 3-5 km (Chernyshev, 1958) or according to other data 5—6 km (Allayarov, 1964). Burrows and shelters. On the west bank of the Caspian Sea dens for kittens are established in openings among the reeds, on dry islets among thickets of blackberry and clematis, and in thorny green hedges. In the Lenkoran lowlands the cat often lives in deserted burrows of badgers, foxes, and porcupines. It lives in burrows in winter also. Statements to the effect that this cat digs its own burrows are unconfirmed. On June 26, 1925 in the vicinity of the city of Makhachkala a nest was found among reeds under a dense oleaster bush. The nearest reeds were pressed to the ground and served as the foundation of the nest. The [nest] platform itself consisted of short grasses and wool. The diameter of the Fig. 178. Thicket of giant plume grass and tamarisk in *“Tigrovaya Balka”’ preserve. Biotope of jungle cat. Tiger, Bukhara deer, wild boar, jackal, and pheasants also live here. Lower Vakhsh, Tadzhikistan. October, 1960. Photograph by A.A. Sludskii. 292 389 foundation of the nest was 100 ст, and of the platform, about 50 cm; the thickness of the nest was 30-35 cm. The nest had been built on a section of ground which was dry at that time of the year. Kittens with their eyes still closed were found inside the nest. A second nest with kittens was found on June 5, 1926 near Kizlyar. It was located on a dense reedy islet (area about a hectare) roughly 400 m from a small creek. The foundation of the nest consisted of reed stalks and the platform of soft grasses and wool. The overall diameter of the nest was 80 cm and of the platform 40—45 cm. Both of these two nests were well concealed (Yu.G. Heptner and Ognev, 1935). In southern Turkmenia newborn kittens have been found lying directly on sand. Other cases of kittens found directly on the ground or under bushes are also known. In Tadzhikistan jungle cats set up nests in fortlike dense tugais or on fallen reed stalks in plume grass thickets. On these stalks, the cat trains dry leaves of reeds and other plants, forming a nest 1.5 to 2.0 m in diameter, covered from all sides with reeds. All three of the nests discovered were so constructed. Instances of cats inhabiting burrows in this area have not been recorded (V.I. Chernyshev, 1948). In Uzbekistan the jungle cat sets up its den in a similar manner in dense reed thickets. In the lower Zeravshan a nest 80 cm in diameter was found among fallen reeds and cattails of the previous years. The platform was covered with reed leaves and wool. From the platform a 2.0 m-long gallery imperceptibly opened into a space [in the reeds]. A second nest in the Amu-Darya delta was on fallen reeds growing in shallow water 10 cm deep 80 cm from the surface (Allayarov, 1964). Near Samarkand, in the Zeravshan valley, a cat occupied the burrow of a porcupine at a distance from the irrigated land. Here there are neither tugais nor dense growths (V.G. Heptner). Daily activity and behavior. The chaus has essentially a nocturnal way of life, setting out to hunt at twilight, but in the warm season of the year it is often active in the morning and during the day. Outside the breeding period in the Trans-Caucasus it spends much of the day hiding in openings among reeds, thickets of thorny bushes, rock crevices, and from time to time in the hollows of trees (Dinnik, 1914); it also rests in burrows. The chaus is not timid and seldom wary. It falls into traps comparatively easily. It is strong and malicious; rare is the dog which can tackle one by itself. While hunting on land the cat moves stealthily, without noise, close to the ground. It stops from time to time, listens, and sometimes raises its head as though scanning the distance. In stalking a quarry it almost creeps along the ground, hunches its shoulders tightly, pricks up its ears, spas- modically switches its tail, sometimes freezes in place, later creeps forward again, and finally makes a decisive leap. Its leaps are not long. If it misses 390 on the first pounce, it makes one ог two more leaps, after which the prey is not pursued. While hunting it is guided not only by audition and vision, but also by olfaction. Repeatedly cats have been observed to sniff constantly as they proceed along a path. Sometimes it bypasses well-hidden traps set in the trail. It often climbs trees; on one occasion a suckling [pig] killed and hung on a tree by hunters was partly devoured by a cat (Dinnik, 1914). Sometimes the cat lies in wait for its prey for quite some time, remaining concealed close to the trail in openings in reeds, and at other such sites. An observer once happened to see one jump from behind a bush on a desert hare running past and quickly dispatch it. Jungle cats usually hunt in the early morning and evening hours and rest at the end of the day. They also hunt in the first half of the night and have quite often been encountered searching for prey even during the day. Injured ducks in particular are frequently hunted during the day. In areas where hunting of game is common (Trans-Caucasus), cats regularly gather from the immediate vicinity. They are not afraid of gunshots, remain close to hunters, and quickly seize undetected wounded birds (Е.Р. Spangenberg). Magpies interfere in day hunts of jungle cats by following the predator and emitting loud alarm calls. Even when caught as a kitten, a jungle cat is quite difficult to tame. One animal, taken as a kitten only 30-40 days old, remained vicious the entire time. No one could extend a hand to it; it growled, attacked with its claws, and inflicted injuries even on a hand covered with a glove. This cat adapted to a diurnal mode of life but was especially active in the evening under artificial lighting. It was trained to jump from one stool to another and to walk a thin pole. While performing these exercises it issued loud calls and threatened the trainer constantly with its claws; it never attempted to bite him, however (N.N. Rukovskii). Seasonal migrations and transgressions. These aspects are not known. Reproduction. In Dagestan and the Trans-Caucasus, judging from the period of appearance of kittens, estrus would seem to be greatly prolonged, extending from the end of January through the first half of April inclusive. Hence statements (Vereshchagin, 1942) that mating in the Caucasus occurs in March and in the eastern Trans-Caucasus in February (oid calendar; Satunin, 1915) are only partly true. In southern Turkmenia mating occurs in January to early February. In southwestern Tadzhikistan the testes greatly increase in weight by January and become particularly large in February- March. The testes of a cat caught in October weighed 1.03 g while those of another animal caught in February weighed 4.62 g. Active spermatogenesis occurs in February and the first half of March but, considering the period of the appearance of kittens, one could assume that it occurs even in January. In the middle of March spermatogenesis slows 293 294 hh down but spermatozoa are preserved in the epididymis until mid-May (two cases). Based on the condition of the testes, occurrence of pregnant females, and periods of birth of kittens, one may assume therefore that the period of estrus usually extends from January to March in the region under con- sideration. In warm winters estrus commences almost one month earlier than in severe winters. In ‘‘Tigrovaya Balka’’ preserve on the lower Vakhsh the earliest mating of these cats was observed on January 10, 1948 and the last on February 15, 1950. It is possible ihat in spring some individual females mate even later. In 1950 two adult females, not yet pregnant, were caught on March 25 and 30 and another nonpregnant female even on April 10 (V.I. Chernyshev). During the period of estrus, after sunset and throughout the night, the shrieks of males are heard, which differ from the normai howls of domestic cats in a more prolonged “‘gurgle’’. Violent combats occur between males in rut as evidenced by traces of blood and tufts of hair. The duration of gestation is not precisely known but evidently covers a period of about 60 days. In Dagestan kittens are seen from the end of March to early June. The earliest kittens have been discovered in the Kyzlyar region was the last few days of March, 1926. Near Makhachkala three kittens with their eyes still closed were discovered on June 20, 1925. Two fairly grown kittens were found near Kyzlyar on June 5, 1926 together with a lactating cat; another very large kitten was found here in the first half of July, 1925 (Yu.G. Heptner, Heptner and Formozov, 1941). In the Trans-Caucasus (Lenkoran region) on June 14, 1933, a litter of three kittens with their eyes open but still weak in their legs was found (E.P. Spangenberg), and another three in the same condition happened on May 25, 1951 at Kyurdamir (Azerbaidzhan). Two females caught at Kyurdamir on June 7 and 10 were found to be lactating and had considerable milk in their glands; four placental scars were found in the uterus of one (Yu.A. Gerasimov). In the Caucasus young are seen in early May (Vereshchagin, 1942). In southern Turkmenia at Saryyaza on March 29, 1887 (old calendar) five newborn kittens were found (Radde, 1889). Kittens with a body length of about 20 cm were found on April 5, 1901 (old calendar) in the Tedzhen valley (Zoological Museum, Academy of Sciences). In southwestern Tad- zhikistan the first pregnant female was caught on February 13, 1948, and contained three fetuses weighing 13.7 to 14.2 g. A second cat caught on February 21, 1950 was found to be in estrus. Of two females caught on March 3, 1949, one had four fetuses aged about 30 days and the other three aged 20 to 25 days. A cat caught on March 21, 1950 had only just mated; three corpora lutea were found in her ovaries. The earliest date of all births recorded was March 31, 1949; this parti- 293 392 So Fig. 179. Impenetrable stronghold of oleaster (E/eagnus), plume grass, reeds, and liana. Residence of jungle cats in Pyandzh valley. Tadzhikistan. April, 1966. Photograph by G.N. Sapozhnikov. cular female produced four kittens. A female caught on April 18, 1950 was already lactating. The last of the births recorded took place on May 10, 1947 (Tadzhikistan, Parkhar region) (Chernyshev, 1958). Judging from the periods of estrus and sightings of pregnant females, kittens in the Parkhar region (Tadzhikistan) may evidently be expected anytime between mid- March to mid-May, possibly even later. The litter consists of three to five young, usually only three (14 obser- 393 vations ). Among cats breeding in Alma-Ata Zoological Garden there were usually three kittens per litter; four were found only once. References to the occurrence of up to 10 kittens in a litter are apparently erroneous (Kashkarov, 1932; Sultanov, 1939; Naumov and Lavrov, 1941). The well-known report of Blanford (1888-1891), quoted by later investigators, mentions that in India this cat annually produces two litters of three to four kittens each. This cat sometimes produces two litters in a year in the Soviet Union also. Thus in Ismaillinsk region of Azerbaidzhan pregnant jungle cats were encountered in summer also. At the end of August, 1952, a nest deep under the roots of a tree contained kittens about 30 to 40 days old (their eyes were open and their teeth erupted) (N.N. Rukovskii). -In the Zoological Museum, Academy of Sciences, there is a kitten weighing one-fourth the wei ght of an adult with the label ‘‘19 October, 1936, western Kopet-Dag, Chandyr River’’. This cat must have been born in August. A pair of jungle cats from southern Turkmenia littered twice in a season in Alma-Ata Zoological Garden, i.e., in March-April and in August; in 1957 they even produced three litters in a single season (the female did not nurse the young). In Uzbekistan in the Angren valley kittens have been seen at the end of April and in early August (Lustin and Putsatov, 1957). In Tadzhikistan a high percentage of barren females has been noted among female jungle cats. Of the 11 adult females caught during the breeding season, only 7 (63%) were pregnant and 4 (37%) barren. The reasons for this phenomenon have not been established (Chernyshev, 1958). Growth, development, and molt. Kittens are born blind, with their ears closed, and helpless. They are covered with a dense furry coat of hair. The general color shade of the juvenile coat on the head and back is a light brown, gradually fading toward the flanks. The underside of the body is light olive. On the forepaws there are four faint transverse bands, and on the rear paws, six. Faintly visible dark-colored spots occur on the flanks and underside of the body. In the rear of the back there are four longitudinal dark bands. Longitudinal bands also occur on the crown and neck. The tail is the same color as the back. Among kittens one-fourth the size of adults, bands are present on the rear paws and flanks. The weight of four two-day-old kittens born on March 31, 1949, was 45, 50, 43 and 55 g. The kittens gain sight on the tenth to the twelfth day. Lactation continues for about two months. In southwestern Tadzhikistan the cessation of lactation in different years varied between June 10 and July 20 (Chernyshev, 1958). Milk teeth were shed by a young captive cat and replaced by permanent ones at the age of five months (N.N. Rukovskii). The litter of this species breaks up early, at the end of summer, and hence the animals are invariably seen singly in autumn and winter. By October-November the young sometimes attain one-half the size of adults 394 and weigh at that time 5.0 to 6.0 kg. In winter they grow very slowly and hence young animals in spring weigh the same as in [the previous] autumn. In southwestern Tadzhikistan young females weighed 2,400 g on November 12, 1949; 2,700 g on March 30, 1950; and 2,618 on April 1, 1950 (the weight of the largest adult female was 7,500 g). A young male caught on March 5, 1947 weighed 5,000 g (the adult weighs about 8,000 g, and even up to 12,000 g) (Chernyshev, 1948). A tame chaus aged 7.5 to 8.0 months weighed 7.0 kg and was not obese (N.N. Rukovskii). The jungle cat molts twice a year. In the Trans-Caucasus spring той. commences in February and ends by mid-April. Cats caught in June sport a summer coat and a clean light-colored inner skin surface. Autumn molt is more protracted. It commences in September and ends in the first few days of November. Only after this period does the skin become completely renewed. Reference to molt occurring in May in the Caucasus is errone- ous (Vereshchagin, 1942). In southwestern Tadzhikistan in warm winters jungle cats begin to molt in the first few days of February. The limbs and heads of specimens caught in the middle and end of February were completely molting. By the end of April all animals have completed the molt. In the cold winter of 1949/1950, when very low temperatures prevailed from December to beyond the first half of February, molt was delayed. Cats caught in early and middle March still bore a winter coat with faint traces of molt. Nevertheless, by early Мау а summer coat was seen in all animals. Thus, in this cold winter the commencement of molt was delayed by one month. In autumn the shedding of summer hair (guard hair) and the growth of underfur were seen from mid-October on. Molt was completed by the end of November to mid-December. The winter coat is more grayish in hue while the summer coat is more sandy, blending well with the surrounding dry vegetation (Chernyshev, 1958). Enemies, diseases, parasites, mortality, competitors, and population dynamics. Enemies and diseases are not well known. Kittens caught in Ismaillinsk region (Azerbaidzhan) were infected with coccidians (N.N. Rukovskii). In Georgia the following helminths were found in chaus: Hydatigera taeniaeformis, Mesocestoides lineatus, Toxocara mystax, Capillaria feliscati, Thominx aerophilus, and Ancylostoma caninum (Rodonaya, 1951). In Azerbaidzhan seven species of parasitic worms have been found: Diphyllobo- thrium erinacei, Hydatigera krepkogorskii, H. taeniaeformis, Mesocestoides lineatus, Troglostrongylus assadovi, Toxocara mystax, and Petrowospirara petrowi (Sadykhoy, 1955). In southwestern Tadzhikistan 10 species of parasitic worms and one parasitic larva have been detected (Chernyshev, 1953) (Table 13). 296 395 Table 13. Helminth parasites т jungle cats of southwest Tadzhikistan (upper Amu-Darya; СБегпу$Веу, 1953) Helminth or its larva Number of Degree of Number of Location of infected infection parasites _ parasite animals (регсеп- tage) Diphyllobothrium mansoni 11 47.3 1-31 Small intestine Sporganum mansoni 2 8.6 2-60 Body cavity, muscles, subcutaneous tissue Hydatigera taeniaeformis о 30.7 2-26 Small intestine Mesocestoides lineatus 4 172 3-27 Large intestine Toxocara mystax 11 473 1-24 Gullet, abdomen. and intestine Uncinaria stenocephala 1 43 ? Large intestine Gnatostoma spinigerum 1 43 6 Abdominal walls and cavity Dirofilaria immitis il 43 1 Pericardium Dirofilaria repens 2 8.6 2-3 Subcutaneous tissue, capsule, kidneys Dracuncultis medinesis 2 8.6 2-7 Subcutaneous tissue Physaloptera praeputialis 3 12.9 1-2 Stomach In Uzbekistan the bronchi of one of eight cats dissected were infected with the nematode Proglostrongylus badanini (Muminov, 1964). The number of helminths in a kidney, usually the right one, can be so high that its weight may exceed the weight of the other by twofold (Reimov and Nuratdinov, 1970; Amu-Darya delta). In southwestern Tadzhikistan, the ticks Rhipice- phalus turanicus, К. sanguines, another nonidentifiable Rhipicephalus species, and Hyailomma anatolicum were found in these cats. Maximum infestation by ticks was observed from May through September (V.I. Chernyshev). In southwestern Tadzhikistan the fleas Pulex irritans and Xenopsylla nesokiae wete found in two animals. The latter flea is a parasite characteristic of the complex-toothed rat. The competitors of jungle cat are jackals, forest and steppe cats, foxes, and predatory birds, especially the marsh harrier [Circus aeruginosus]. The main competitors are jackals and forest cats, which feed on almost the same animals. Their importance as competitors can be judged from the following: in the Trans-Caucasus, near Shil’yan, 290 jackals and 68 jungle cats were caught around a group of lakes in the winter of 1947/1948. Usually jungle and forest cats do not live in the same [geographic] place or biotope. If, however, they occur in the same region, the one species is always the only 396 one. It is significant that jungle cats are absent on the Black Sea coast of the Caucasus and in the swamps of the Kuban, where forest cats are abundant or common. Judging from the record of the tanned skins of chaus in the southern part of its range, its population fluctuates from year to year, but not much, deviating from the multiyear mean by not more than 30%. Factors responsible for fluctuations in population numbers have not been studied. In the northern part of the range fluctuations in population numbers are evidently due to extreme cold and abundantly snowy winters, when all water sources freeze and deep snow cover prevails for a prolonged period. In the southern part of the range, however, these very same conditions are distinctly favorable for the cat; in such winters it enjoys an abundance of food in the form of aquatic and other birds weakened by lack of food. In some years, however, the southern population may undergo a reduction due to epizootic diseases (carnivore distemper, pasteurellosis, paratyphoid, rabies, and others) (also see ‘‘Population’’). Field characteristics. A large cat (weighing up to 12 kg), standing tall on its legs, with a monochromatic reddish-brown coat. The tail is rather short and thin and the ears erect with tassels. Usually, it is seen running in an unhurried, slow gallop. While running, it sways slightly from side to side. It is often encountered near water, in openings in reeds, in kupak*, or while swimming. Its tracks are often seen on sandbars, spits, and silty banks and are much larger than the prints of domestic, steppe and forest cats. The imprint of the paw of a jungle cat is 5.0 cm x 6.0 cm in size. Moreover, the digital impressions are widely separated, while in other cats they are compact. The impression of the hind sole is trapezoidal and not rounded as in the forest cat (Fig. 180). Tracks extend in the form of zigzag line. The pace length is 29 to 32 cm. The meow of a jungle cat is similar to that of domestic cat but deeper. The voice is heard rather frequently at night, especially at the end of winter and in spring. (A.S.) Practical Significance Jungle cats are fur-bearing animals but their pelt are of little value. The All- Union standards include their pelts in the group ‘‘wildcats,’’ in which they are distinguished as ‘‘jungle cats (sabanchi)’’. Commercial hunting of the jungle cat was practiced even in the eighteenth century. P.I. Rychkov (1762) wrote: “‘Wildcats were present *Floating reed mats—Sci. Ed. 296 397 Fig. 180. Tracks of a jungle cat on silt. ‘“Tigrovaya Balka’’ preserve at Vakhsh. Tadzhikistan. October, 1960. Photograph by A.A. Sludskii. among Kirgizian hordes on the steppes, but few brought in for sale by the Kirgiz-Kaisak in Orenburg and Troitsk Fort. They were far bigger than domestic cats and resembled lynx, though smaller, and the fur was poorer. A single animal sold for 16 to 20 kopecks.’’ Judging from the description of the skin these were jungle cats. At present, in fur-tanning factories this cat is of some importance only in Azerbaidzhan, where 2,966 to 4,300 or an average of 3,500 skins were tanned annually from 1931 through 1940 (these also include pelts of forest wildcat) (Vereshchagin, 1942). In Tadzhikistan in 1946 about 300 skins were prepared (Chernyshev, 1948). In Turkmenia the tanning of pelts of jungle cat has failen considerably in recent years, mainly because of the absence of commercial hunting. Here 91 animals were caught in 1954, 109 in 1955, 187 in 1956, 172 in 1957, and 166 in 1958. There is no information for the rest of the Union Republics since jungle cat is reckoned as *‘wildcat’’ by fur-tanning factories. This cat everywhere is a serious pest of the hunting economy. All investigators regard it as the main enemy of pheasants, francolins, and 398 waterfowl. At several places, although the francolin was protected from hunting, its population did not rise due to its destruction by the jungle cat (Satunin, 1915). Like the wolf, this predator greatly damages the nutria farming of a semiwild type. Next to and within the precincts of a nutria farm about 200 cats were caught in traps over a period of 14 years. In Kyurdamir (Azerbaidzhan) since 1946, a bounty of 5 rubles is paid by nutria farms for the skin of a cat. At places, especially in the Amu-Darya delta, the cat adversely affects the muskrat economy. In some regions this cat is also a pest of poultry farms. No special hunting is organized to catch this cat. It is caught incident- ally during hunts for pheasants and pigs; it also falls into traps set for fox or jackal. It can be caught comparatively easily in a trap placed in water at a shallow place with a live fish fastened to a rod run through its gills. The splashing of the struggling fish attracts the attention of the - cat (Yu.A. Gerasimov). Pelts are used in making ladies’ fur coats or imitation sealskin. In view of the harm inflicted by this predator, its capture is permitted everywhere year-round. In Azerbaidzhan a bounty of 5 to 12 rubles is paid for each skin brought in. In old literature there exist references that in ancient Egypt, judging from mummies found, the jungle cat was domesticated; but later it was not found in a domesticated state (Bogolyubskii, 1959). Ancient Egyptians successfully crossed domestic cats with chaus and mummies of such hybrids have purportedly been found. It has also been reported that hybrids were used in hunting waterfow] and for tracking wounded birds; some pictures have been preserved depicting various scenes of such hunts with hybrid | cats (Bogdanov, 1913). These data have found no confirmation in recent literature (see the section on the origin of domestic cats under group /ibyca and ‘‘Systematic Position’’ of jungle cat). (A.S.) WILDCAT Felis (Felis) silvestris Schreber, 1777 1777. Felis (Catus) silvestris. Schreber. Sdugeth., vol. 3, р. 397. Germany or northern France. In the species F. (F.) silvestris are here included two groups of forms earlier considered independent but closely related species: Р. (F.) silvestris Schreber, 1777—forest cats of Europe, Asia Minor, and the Caucasus, and Е. (F.) libyca Forster, 1780—steppe (spotted, bay, and long-tailed) cats of Africa and Asia. The basis for this unification was indicated in the last century but 299 399 formalized only in the present-day (Haltenorth, 1953). Both species are morphologically very similar and there are no constant craniological differences between them in the various parts of their range (Pocock, 1951).27 Differences between the two species are confined to coloration and some features of the pelage (general characteristics of fur and fluffi- ness of tail). Within the limits of each ‘““species’’ geographic variation does occur but nonetheless the distinctive features of both are linked by intermediates. Both ‘‘species’’ and all their geographic forms are strictly vicarious. All forms belonging to the European forest cat group (si/vestris), like those of the steppe cat group (ПБуса), hybridize in nature with the domestic cat, which itself is a domesticated steppe (‘‘Ъау’”) cat of northeastern Africa, Е. 5. libyca (see р. 452). In some regions of Europe it is believed that the entire population of wildcats has some, even significant, admixture of the blood of domestic cats, and that evidently the hybrids may be regarded as fully fertile (this has also been established experimentally). Domestic cats, especially in countries with snowfree winters or those with little snow, readily turn feral, and some populations asserted or suggested to be separate species or subspecies merely represent feral domestic cats (megalotis—Timor, agria—Crete, daemon—Caucasus, mediterranea—ltaly, pulchella— Africa, syriaca—Syria, and issikkulensis—Semirech’e). The range of the species silvestris, as interpreted here, is very extensive (Fig. 181). However, it is unified and quite natural. The range of this cat is quite similar, broadly speaking, to the range of other widely distributed cats of the Old World (lion, leopard, caracal, and cheetah). The range occupies Europe, including England in the west (absent in Ireland and never present), the Baltic region in the north, and up to the Dnepr and Don Rivers in the east; Africa except the dense humid forests of western Africa; the Near East, including Asia Minor, the Caucasus; northwestern India; Middle Asia and southern Kazakhstan, Dzhungaria, Kashgaria, Inner Mongolia, and southern parts of Mongolian People’s Republic (reconstructed range). The entire mainland of Europe and England are inhabited by typical members of the silvestris group. The same is true of the Caucasus and Asia Minor, at least over much of it. Both in the Caucasus and in southern Asia Minor, the ranges of subspecies of the silvestris approach very close to, and adjoin at places, the ranges of subspecies of the /ibyca group. Transitional populations however are absent; and hence transitional animals are not known there. "7Craniological differences detected between Turkestan steppe and European (Caucasian) forest cats (V.G. Heptner; see pp. 75 and below), not observed by earlier authors, could not be verified in all the geographic forms. 400 4 J \2 | | Ny ] | i | at 298 1000 0 1000 2000 3000 4000kin a a ee 90 Fig. 181. Reconstructed species ranges of forest and steppe wildcat, Felis (Felis) silvestris Schreb. s. |. (schematic) (scale in km). Region of occurrence of European forest cat—subspecies group silvestris (in the northwest), and steppe cat— subspecies group /ibyca (rest of the range) separated by broken line. Mediterranean insular transitional forms are included in the European forest group. The northern boundary of the range in Kazakhstan is tentative (see Figs. 198 and 267). (V.G. Heptner). The Mediterranean island forms and in part those of northern Africa [jordansi, reyi, cretensis (agria), and sarda] represent intermediates. Whatever be the characteristics of these individual forms and their history (degree of admixture of blood of domestic cats, domestic cat simply turned feral, etc.), some of their features are a mixture of features of European forms of the silvestris group and of the African libyca group. In the nature of 300 401 their fur and color and fluffiness of tail, they belong to the libyca type, but the skull shows features of the silvestris type.* Even the diversity in their interpretation underscores their unique (intermediate) taxonomic position. At the beginning of the century (Miller, 1912) they were considered completely independent species (sarda and agria-cretensis) but nowadays all have been combined into one subspecies (sarda); some taxonomists place the subspecies in species libyca (Pocock, 1951; Ellerman and Morrison-Scott, 1951 and 1966), while other maintain that they occupy an intermediate position between the “‘species’’ silvestris and /ibyca (Haltenorth, 1953 and 1957). This latter view is very widespread. The systematic relations of the two groups of forms viewed from the process of speciation are of extreme theoretical interest. They may be assigned an intermediate position between species and subspecies or the entire complex of forms considered ‘‘superspecies’’ or ‘‘macrospecies’’. A description of the species, including synonyms, has been given separately for the two ‘‘groups’’ of subspecies: European wild forest cats, Е. (F.) s.—‘‘silvestris group’’; and steppe wildcats, F. (F.) /.—‘‘libyca group’’. Only in the sections ‘‘Diagnosis”’ and ‘Systematic Position’’ have the two been presented together. (V.H.) Diagnosis Size small; like that of domestic cats or only slightly larger. Background color gray of different intensities, being very light-colored in some forms with a fairly intense ocherous shade. Dark spots are present, or a dark belt and spots on the back, or a belt and transverse stripes on the trunk, or a belt without distinct spots or transverse bands. Dark longitudinal bands occur along the top of the head. The tail has dark spots or rings (transverse). The coat is thick and rich with dense underfur, or relatively short with slightly developed and close-fitting underfur. Tail furry, thick, and blunt, or thin and pointed at the end. Skull not elongate, and zygomatic arches wide-set. Palate not extended into interpterygoid vacuity posteriorly (see Fig. 28) and interpterygoid vacuity broad. Second premolar (first in row) generally present; antero-inter cusp of upper carnassial tooth well developed and its peak distinct but small and blunt. Ectotympanic chamber of auditory bulla poorly developed (see Fig. 24). Maximum diameter of auditory opening not more than length of third (second in row) upper premolar and part of auditory bulla between auditory 8One of the characteristics of these forms, judged from the limited amount of material from Sardinia (sarda) available in the Zoological Museum, Moscow University, is the similarity in structure of auditory bulla to that of silvestris forms, i.e., the small bulge in front of the ear canal in the ectotympanic part is absent (see Fig. 29). 301 402 and Eustachian openings does not bulge (European wildcat group, si/vestris); or maximum diameter of the external auditory meatus more than length of third upper molar and auditory bulla between auditory and Eustachian openings somewhat inflated and shifted forward slightly more than portion lying inward from Eustachian tube (steppe cat group, libyca; Fig. 29). (V.H.) A. GROUP SILVESTRIS—EUROPEAN FOREST WILDCATS 1777. Felis (Catus) silvestris. Schreber. Sdugeth., vol. 3, p. 397. Germany or northern France. 1777. Felis catus ferus. Erxleben. Syst. Regni Anim., vol. 1, p. 518. Southern Europe. 1837. Felis cato affinis. Hohenacher. Bull. Soc., Naturalistes Moscou, 1, 7, 136. Elenendorf, Elisavetpol’ governance (now Khallar, south of Kirovabad, Azerbaidzhan). 1896. Catus ferox. Martorelli. Atti Soc. Ital. Sci. Nat. Milano, vol. 35, p. 275. Thuringia. This name arose through some misunderstanding (described together with Catus ferus Brehm, 1869) but was never- theless used in the nomenclature for this form in the Russian fauna. 1904. Felis daemon.” Satunin. Proc. Zool. Soc. London, p. 162, Elisavet- pol’ (now Kirovabad, Azerbaidzhan). 1905. Felis catus caucasicus. Satunin. Izv. Kavk. Muzeya, Mitt. Kauk. Mus., 2, 2-4, 154. Borzhom. 1916. Felis silvestris trapezia. Blackler. Ann. Mag. Nat. Hist., vol. 18, p. 73. Khotz near Trapezund [Trabzon], Asia Minor. 1943. Felis silvestris euxina. Pocock. Ann. Mag. Nat. Hist., vol. 10, p. 701. Baspunar in Dabruja, Romania. (V.H.) Description Size small, but averaging larger than that of domestic cats. In general appearance the forest cat is very similar to the domestic cat —body long and stands on rather short legs, and tail comparatively long (Fig. 182); its length usually slightly exceeds one-half of body length (sometimes comprising 70% of it), or equal to it, and more rarely shorter. At the same time, partly due to the generally greater size, but mainly because of the much longer and richer coat, especially the winter coat, the forest cat appears far bigger, powerful, and more massive than the domestic cat. Head broad; jugal region with short, blunt maxillary portion seemingly *°Black feral domestic cat or melanistic forest cat. 301 403 Fig. 182. European wildcat. Berlin Zoological Garden. Photograph by С. Ви КВ. less extended—skull more spherical than in jungle and Amur cats. Ears moderate in length, quite wide-set, and without tufts at tip. Hair at tip nevertheless may be somewhat long—up to 3.0 to 4.0 mm. Ears in winter project out of fur. Side whiskers absent on cheeks. Vibrissae white; hairs on lips range from 7 to 16 on each side and reach 50 to 80 mm in length, eyelashes number 6 to 8 each and range from 50 to 60 mm in length; group of 3 to 6 vibrissae 30 to 40 mm in length present on inside of wrist (Haltenorth, 1957). Eyes large, pupil vertical, and iris yellowish-green. Bare region on nose in living animal flesh-colored (not black). Pelage throughout body fairly uniform in length (long hair absent on neck or back). Hair on tail very long and dense and hence tail appears furry and thick. At end of tail hair not shortened and tips not pointed but thick as Ww 404 though chopped (characteristic feature of forest wildcat). Length of guard hair in winter coat 70 to 72 mm, top hair 55 to 60 mm, and underfur 45 to 55 mm; corresponding measurements in summer coat 50 to 67, 45 to 60, and up to 53 mm. Thickness of top hair 90 to 100 microns and of underfur 20 microns. On the average 10 to 20 guard hairs, 65 to 100 top hairs, and 1,600 to 2,100 underfur hairs occur in an area 25 mm? of the winter coat along the middle of the back; corresponding figures for the summer coat are 6 to 9, 66 to 100, and 380 to 1,100 (central European form; Haltenorth, 1957) (see ‘‘Geographic Variation’’ for description of pelage of Caucasian cat). Main color shade of winter coat fairly light, gray, somewhat richer along the back and faded toward the undersides of the flanks.*° Ocherous bloom imperceptible or barely discernible only along underside of flanks. Very distinct, fairly narrow black band commences at the shoulders, runs along the back and usually terminates slightly short of tail base; however, it sometimes reaches the tail or even extends onto it as a narrower band along the dorsal surface. Color on flanks monochromatic; transverse stripes from dorsal band either absent or present only as hazy, indistinct shadows; they are rarely distinguishable as stripes and better identified as indistinct dark smudges; sometimes, especially on underside of flanks, they have a rusty tone. Fairly distinct but not very sharp transverse stripes, not more than five to seven, are a rare occurrence. Transverse bands on the legs are either vague or undeveloped and only occasionally fairly distinct. Undersurface of body covered with very light gray hair with a light ocherous tinge; dark spots absent or occasionally a few present. On the throat a white spot may be present (quite rarely) or three such spots occur — one on the throat, another between the forelegs, and a third in the rear (inguinal) region of the abdomen. Posteriorly the abdomen is generally a pure and quite bright ocherous color. Color of tail same as on the back; tip of tail pure black. Above tail tip usually two, more rarely three clear, black, transverse rings occur, which are narrower than the black tip and separated from it and from each other by light-colored sections; width of sections roughly equal to the width of the rings. Usually, in front of the black rings, two or three more indistinct, rather dull, sometimes barely perceptible dark-colored, most often brown rings occur. Sometimes along crest of tail there lies a narrow dark band, which is a continuation of the band along the spine that joins the rings on the tail. This pattern is found in cases where the number of rings is greater *Description based on a series of Caucasian skins available in the Zoological Museum, Moscow University. This description differs significantly from that given by S.I. Ognev (1935) since the latter had only limited material available to him and that, too, not wholly representative but with a relatively well-developed pattern. ‘лолешоу ‘КУ Aq Зицатеа ‘итипеб DoISvONDD 514452115 (51251) SYA ‘eo 38910} UBISEONeD “¢ за. 303 405 and their color brighter, and general color of the tail also more vivid. Main color of lower part of the tail slightly lighter than upper part. Upper surface of front paws monochromatic, grayish-yellow, and same color as foot but more distinctly rust-toned. On the lower surface of the foot, in the anterior part, a dirty blackish-brown section covers one-third or one-half the area of the paw (remainder dirty ocherous-gray) or sometimes the whole of it.>! Dorsal surface of the neck and head same color as dorsal surface of the trunk, but around the eyes, lips, cheeks, and chin color lighter gray with a fair admixture of ochre. Hair on inner surface of ear yellowish-white and also forms a sort of border on outer surface. Outer surface of ears dark gray with a rusty or ocherous tone. Forehead and top of head bear pattern of four well-developed black or brownish-black bands; sometimes these bands split, on forehead, into small spots extending onto neck. In the shoulder region, in front of black dorsal band, usually two short, narrow, dark stripes present. From outer corner of the eye, under ear, and beyond onto side of neck, runs a dark narrow stripe; another similar stripe commences under the eye, slightly away from it, and also extends onto neck. Summer coat with fairly light, pure background color, without admixture of ocher or brown; it is even ashen-colored in some animals. Pattern on back and head well developed like pattern on tail, but pattern on flanks very indistinct and may be altogether absent. There are no sex-related differences in color although possibly females may be somewhat more gray than males (Dinnik, 1914). Pelage of young animals at birth has a very light general background with very vivid velvety black spots. Bands and spots numerous, sharply defined, vivid, and cover greater area than general background color. Spots in adult animals possibly fainter than in old animals. Geographic variation in color is not pronounced. The intensity of general coat color and number of spots and stripes and their degree of vividness vary moderately. On the whole coloration conforms to Gloger’s rule, with greater vividness and more prominent spottiness in forms from Scotland, and an intermediate coloration evident in central European forms. Preanal glands present in both sexes; moderate-sized sweat and sebace- ous glands are located around the anal opening and larger-sized sebaceous glands and a few scent glands dispersed in the caudal region, extending along the entire tail on dorsal side. In males, preanal pockets are present in the З!ТВе dark field, when it covers the entire undersurface of the paw, is usually considered (Suminskii, 1962; see later) due to an admixture of blood of domestic cat. Among Caucasian cats this occurs merely as an individual variation. 406 / May aa и cs A \ gi SY NSA on \ А 3 y wid Ve hy , 303 Fig. 183. Skull of Caucasian forest wildcat, Felis (Felis) silvestris caucasica Satunin. No. S 14230, collection of the Zoological Museum, Moscow University, old male. Near Tarsk station close to Ordzhonikidze. December 6, 1928. Sketch by N.N. Kondakov. form of two folds on each side of the anal opening with channels for 304 discharging secretion. These pockets, like the glandular area along the dorsal 407 surface of the tail, are activated with the onset of sexual maturation and play a significant role in reproduction and marking of territory (Haltenorth, 1957; for domestic cat). Teats, four pairs —two thoracic and two abdominal. The skull of the forest wildcat (Fig. 183) differs significantly from that of the jungle cat in somewhat smaller size and weight (for the same age). It is significantly less extended and the zygomatic arches more wide-set not only medially (zygomatic width three-fourths of condylobasal length), but also anteriorly, where they are more steeply set in relation to the sagittal plane. General outline of skull also more spherical since the cranial region is not so constricted and extended posteriorly. Upper line of profile bulges while the large frontal area bulges only slightly, or is almost flat, or exhibits a very gentle and barely perceptible depression. Orbits relatively large, quite rounded, and set forward to a greater degree. Supraorbital processes project perceptibly downward and often are so well developed that they almost meet postorbital processes of jugal, or even fuse with them to close the orbits from behind. In front of orbits, maxillary (facial) section of skull highly compressed and jaws therefore parallel. Nasals generally terminate posteriorly quite bluntly, but in some individuals rather sharply; distance between anterior processes of frontals and upper ends of premaxillae greater. Lateral margins of nasals highly compressed medially and relatively broader anteriorly than in jungle cat. Sagittal crest in old animals well developed but only in posterior part of parietals; lambdoidal crests fully developed, and relatively stronger. Auditory bulla occupies roughly the same area as in jungle cat but smaller in size since less inflated, especially in ventral and anterior directions. Anterior margin of bulla does not reach level of glenoid fossa and postglenoid process. Maximum diameter of opening of auditory meatus less than length of upper third premolar (anterior to carnassial tooth) or equal to it. Bulla not inflated in front of opening of auditory meatus and chambers on either side of Eustachian opening roughly equal in size and lie at approximately the same level.>? Palate not extended posteriorly into interpterygoid vacuity; its posterior boundary either lies at level of posterior edge of molars or only slightly projects beyond that level. Interpterygoid vacuity relatively narrower than in jungle cat and more uniform in width throughout its extension. Posterior edge of palate with small medial tapered projection. Palate itself relatively shorter than in jungle cat and somewhat relatively broader posteriorly. In conformity with more sharply sideward-set zygomatic processes of maxillae, tooth rows do not form a straight line from first Зее notes about this characteristic under description of steppe cat. 305 408 maxillary tooth to axis of carnassial as in jungle cat; instead this line is somewhat arced posteriorly. Dentition complete, 1.е., second upper premolar (first in row of maxillary teeth) present and functional. However, this tooth is frequently absent. Thus, they were absent on both sides in 6 (35%) out of 17 males examined from the Caucasus. Among all but 2 skulls of 23 females the second premolar was absent, and seen only on one side in 2 animals. By and large dentition, especially of the maxillary teeth, is far weaker than in jungle cat—not only in length of tooth but also in size. Anterior inner cusp of upper carnassial tooth much smaller than in jungle cat and summit far less developed and blunter. Canine relatively small but quite powerful and long. Skull of female characterized by smaller size, slightly lighter overall build, relatively less intense postorbital constriction, relatively close proximity of semicircular lines (impressions of masseter muscles on roof of skull), and characteristics of dentition already described. Age-related changes seen in intense swelling of brain case in young animals, relatively short facial ‚рай of skull, relatively broad interorbital and especially postorbital constrictions, absence of crests, and wide distance between temporal lines of skull. Geographic variation of skull apparently does not affect its structural features, but some insignificant differences in overall size are known. Thoracic vertebrae number 13, lumbar 7, sacral 3, and caudal 17-23. Length of os penis in adults 5.0 to 6.0 mm and its width 1.0 to 3.0 mm (Haltenorth, 1957). Weight of heart in young male (December) 10 g and heart index 4.0%o; corresponding values for an adult male (January) 24.5 © and 4.36%o. Absolute length of intestine in relation to body length in these animals 1,340 cm and 1 : 2.43 and 1,500 cm and 1 : 2.42 (Caucasian preserve; based on data of Kotov and Ryabov, 1963). Diploid number of chromosomes 38. Body length of males 430-910 mm and of females 400—770 mm; tail length in males 230-405 mm and in females 180-350 mm; length of hind foot 120-160 mm and height of ears about 60-70 mm.*4 Greatest length of skull in males 96.0-112.3 mm and in females 87.0-106.6 mm; condylobasal length in males 86.0—102.6 mm and in females 80-100 mm; zygomatic width in males 62.4-80.0 mm and in females 60.6—69.0 mm; interorbital width in males 16.5—22.0 mm and in females 16.0-21.5 mm; postorbital width in males 31.1-36.0 mm and in females *3Based on 221 animals from various parts of the range of the group, mainly from western Europe, including 9 from the Caucasus and 29 from the Ukraine (evidently, western); males 137 and females 84 (Suminskii, 1962). Some of the low values evidently pertain to juvenile animals. 306 409 30.0-36.5 mm; and length of upper tooth row in males 30.5-33.8 mm and in females 26.7-33.8 тт.“ Data in literature on weight conflict and are often highly exaggerated. The weight of males (185) in Europe reportedly ranges from 3.5 to 15.0 kg and of females (55) from 3.0 to 10.0 kg (Suminskii, 1962). It has even been stated that the weight can exceed these values, reaching 18 kg. The source of information about such gigantic animals is the usual hunter’s exaggerations being published in journals and conscientiously collected by compilers. The existence of very large specimens, perhaps heterotic hybrids with domestic cat, is not denied, but none attains such sizes. No type of food can possibly fatten a cat to a weight of 15 kg. It is significant that in these same publications such divergences and exaggerations are absent for skull sizes and even body measurements. The normal weight of adult males is 5.0 to 6.0 or 7.0 kg; females are somewhat lighter. Cats weighing 8.0 kg are quite rare in the Caucasus (Dinnik, 1914). In Moldavia cats weigh up to.8.0 kg with a body length of up to 830 mm; females weigh up to 6.0 to 6.5 kg and have a body length of up to 750 mm (Lozan and Korchmar’, 1965). Weight varies perceptibly with age, season of the year, and vagaries of environment; when rodents are plentiful, cats are better fed and their average weight higher while, given poor conditions, cats can become severely emaciated (for more details see ‘‘Geographic Variation’’). Some geographic variation does occur in measurements and weight but it is not significant. (V.H.) Systematic Position See above, under notes on the genus on p. 398, and below in the section pertaining to steppe cats (p. 455). Hybridization of Wild with Domestic Cats, and Hybrid Populations Reports exist about the hybridization in nature of forest wildcat and domestic cat over much of the range of the former. This problem, of significant the- oretical interest, is also of practical importance; distinguishing pure and hybrid individuals. The forest wildcat group can be considered forms which evolved autochthonously in Europe during the Early to Middle Pleistocene. The ancestral form was F. /unensis Martelli (Villafranchian; Kurten, 1965). The domestic cat itself represents the domesticated so-called bay cat of Egypt, 34Animals from the Caucasus (Zoological Museum, Moscow University) with the addition of some extreme values taken from European literature (Ognev, 1935; Haltenorth, 1957; Suminskti, 1962; and others). 410 а subspecies of another group of forms of the species (/ibyca, 1.е., F. 5. libyca). The forest wildcat of Europe (group si/vestris) is not in the direct line of evolution of domestic cat (see ‘‘Origin of Domestic Cats’’ under “В. Group ПБуса”). The first appearance of domestic cat in Europe was recorded in the fourth century A.D. (Rome). For a long time this animal was a rarity, but gradually it colonized all Europe (tenth century in England and twelfth century in Kiev; Pidoplichko, 1956). There is no doubt that throughout the Middle Ages and for much of the later period, in spite of intensive felling of forests (‘‘epoch of forest clearing’’ in western and central Europe in the eleventh and twelfth centuries), conditions for the existence of forest wildcat were totally favorable everywhere and the population structure was normal. For these reasons, and also the small size of the population of domestic cat, hybridization with the latter was not significant. Rapid deforestation of western and central Europe and afforestation, especially in the nineteenth century, coincided with the intensification of pursuit and killing of predators (wolf, lynx, and bear) in the interests of promoting game. Wildcats were also victims of extermination and their range shrank rapidly and their population dropped dramatically. Almost throughout the territory of western Europe, apart from some eastern regions (Carpathians), these animals had disappeared or reduced to rare (extremely rare) animals by the early part of the present century. Under these conditions (“‘lack of partners’’ in particular), some zoolo- gists feel that hybridization with domestic cat, stragglers, or even those which had turned feral, became a widespread phenomenon, instead of occasional as at the beginning of the prior century. Several zoologists have suggested the possibility of spontaneous hybridization. According to another viewpoint (Suminskii, 1962 and 1962a) in central and western Europe hybridization was so extensive that, at present, almost all forest wildcats there exhibit features of the domestic cat to some extent or the ‘‘pure’’ forest wildcat is in general absent in Europe. In any case there are no “‘риге’’ populations, although the level of admixture of the blood of domestic cat in different populations obviously varies — еуеп to the extreme extent of predominance of features of domestic cat. Thus the average ‘‘percentage of purity’’*° based on craniological features in the French and Swiss Alps is 44.1%, 5According to this author the degree of ‘‘purity’’ is determined from several features (he suggests more than ten), each of which has been assigned a range of values on a point scale. Their total in a given skull and skin determines the degree of *‘purity’’ of the individual with respect to the forest wildcat. The average of these values provides an idea of the **purity”’ of the population. Suminskii’s scheme is far from convincing since the selection of features is largely arbitrary, all have been assigned equal value, and the range of their individual and age-related changes has not been established (for a detailed critique of Suminskii’s scheme, see Sladek, 1966, and especially Kratochwil and Kratochwil, 1970). 307 411 Hungary 61.6%, former territory of Germany 63.1%, Scotland 66.1%, and Poland 73.0% (Suminskii, 1962 and 1962a). Since the fertility of hybrids is considered low, the destruction of wildcats in Europe has been proceeding not only as a result of their pursuit and ‘‘depriviation of landscapes’’, but also of hybridization. Thus, in spite of measures taken for wildcat protection in some countries, these animals are becoming extinct in Europe, although individual ‘‘wildcats’’ have continued to survive and could possibly build up the population under proper conservation laws. The above situation prevailed more or less in western Europe (see further) but not in the Soviet Union. Conditions for the survival of the animal and its population density in the western regions of the Ukraine, particularly in the extensive Carpathian forests, are such that they enabled the population to preserve a high degree of purity, although instances of hybridization are not excluded. This is even more applicable to the Caucasus. At some places there are reports of domestic cats straying or turning feral but offspring of wild and domestic cats have mainly been observed in small remote forest villages, for example in forest cordons.*° However, the regions of extensive forests in the Great Caucasus and some parts of the Trans-Caucasus are such that hybridization there was practically absent or only a rare occurrence. In any case it could not have influenced the character of the wild population or its purity.*’ While evaluating this phenomenon it should also be kept in mind that the very possibility of female mating with domestic males under conditions of normal density and structure of population of wildcats, as prevailing in the Soviet Union, is perhaps close to nil. Domestic males undoubtedly cannot successfully compete with wild males. Such hybridization can affect the type of domestic cat, which evidently did occur (see below), but not the population type of the wildcat. In the light of recent data the actual position in western and central Europe is not exactly that outlined on the basis of the material studied. It *°Thus for the Caucasian preserve, V.A. Kotov and L.S. Ryabov (1963) have shown ‘innumerable instances of mating (of wildcat) with domestic cats. We have seen many such hybrids in forest cordons . . . . Such kittens exhibit the body form, coloration, and habits of the wild parent. They take fright at the slightest provocation, do not pemnit fondling, and hide the moment strangers appear. As they grow up, such hybrids usually attack domestic birds.”’ 3’Suminskii’s statement (1962) that, according to external characters the ‘‘percentage of purity’ of Ukrainian cats (evidently from western regions and the Carpathians; 29 animals) was 51.6% and that of Caucasian (9 animals) 55.7%, does not appear correct in the light of the foregoing discussion. The discrepancy is underscored by the fact that this index was 50.7% for the former territory of Germany, 45.5% for France (16 animals) and 48.7% for Poland (18 animals). In these countries hybridization is, of course, far more probable. Such contradictions in data underscore either a deficiency in the method itself or incorrect selection and evaluation of indicator characters. 308 412 appears that in spite of all the unreliable distinguishing features and their combinations reported by various authors, forest wildcats and mixed-breed domestic cats (wholly domestic, wanderers, ferals, and solitary) do reliably differ in volume of brain case, 1.е., in size of brain. This characteristic has been detected on the basis of the principle established long ago; namely, that the brain of domestic cat is invariably smaller than that of its wild ancestor (Klatt, 1912). Although the domestic cat has not descended directly from the European wildcat, both belong to the same species. This relationship is maintained in spite of the fact that the cat has undergone less changes during domestication than other species.*8 In the forest wildcat the volume of the brain case exceeds 35 cm3, while it is less than 32 cm? in the domestic cat. When the volume varies between 32 cm? and 35 cm? (rare specimens), these two forms are distin- guished by the ratio of the ‘“‘maximum skull length to the volume of the brain case.”’ If this index is less than 2.75 the skull belongs to a wild cat; if greater, the skull is that of a domestic cat (index true in 99% of 139 skulls of wild and 317 domestic cats recovered from France, former German territory, Romania, Yugoslavia, Hungary and Switzerland; Schauenberg, 1969). This critical index for the west Carpathian population could be reduced to 2.60, while the lower limit of the skull volume of forest wildcats is taken as 36 сш?. In all skulls with a volume index characteristic of domestic cats the depression (glabella) at the juncture of the nasals and frontals is very prominent. This is a characteristic feature of domestic cats (50 skulls of wildcats from western Carpathians and 63 skulls of domestic cats which had become feral; central Moravia-Czechoslovakia; Kratochwil and Kratochwil, 1970). Thus it is evident that the European cat populations under study are essentially pure. This does not exclude the occasional appearance of indi- viduals with some amount of admixture of blood of domestic cats. However, it cannot be said that hybrid populations and the “‘disappearance’’ of wildcats in Europe occurred in this manner. Such an assumption is even less relevant to populations of the Soviet Union, although the appearance of some hybrid animals, primarily perhaps in Moldavia, cannot be ruled out. However, this problem should be resolved in each individual case, apart from obvious ones (“‘domestic’’ color, etc.), based on the volume index of the brain case. Hybrids of wildcats (Е. 5. silvestris and F. 5. caucasica) and steppe cats (F. s. caudata) have evidently not been found in nature. The reason perhaps lies in the absence of extensive and close geographic and biotopic contact “Features suggested by various zoologists for differentiating these two forms of cats, in addition to other details, apart from articles by Suminskii cited in the text, can be verified in works by Ognev, 1935; Вбрт$, 1955; Haltenorth, 1957; and Sladek, 1966. 413 between them. In the Soviet Union such hybrids have not been reported and apparently do not occur.*? (V.H.) Geographic Distribution Found in the forest regions of central and southern Europe, Asia Minor, and the Caucasus. Geographic Range in the Soviet Union The range in the Soviet Union (reconstructed) represents the eastern and southeastern segment of the range of the group. It occupies the western part of the country and is divided into two sections—European and Caucasian (Fig. 184), which are continuous in the south outside the borders of the Soviet Union. The following were the boundaries of the European part of the range in the period of its maximum development over the historic past. In the northwest the range covered Lithuania and southern Latvia roughly to the latitude of Riga, and northeastern Belorussia to the upper Dnepr in the Mogilev district, but apparently a little to the east (former Mogilev gover- nance). Farther on the boundary turned southeast, encompassing part of former Kaluga governance, former Voronezh governance, and in particular Shipov forest on the Don at Pavlovsk. From there the boundary evidently proceeded southwest toward the great bend of the Dnepr, extending south of Zaporozh’, and descended to the sea along the Dnepr. To the west of this line the range everywhere extended to the western Soviet borders and into central and southeastern Europe. For the region between the above line and the Dnepr, information is available about the occurrence of this group at Kursk and Belgorod (Kursk- Belgorod territory, land of Belgorod polko), Putivl’, Sumy Akhtyrka (southern Khar’kov), and along both banks of the Dnepr in the land of Zaporozh’e Sechi. Information for these sites pertains to the seventeenth and eighteenth centuries and only to part of the nineteenth. It is quite possible that some references may have been based on encounters with feral cats,*° but some definitely pertain to wildcats. It should be kept in mind that °Haltenorth (1953), citing Ognev (1930), who cited Smirnov (1922) affirms instances of hybridization of wild and steppe cats in the Trans-Caucasus. This is erroneous; both Ognev and Smirnov mention only hybrids of domestic and steppe cats. 4°Such, for example, was the report pertaining to the ‘former Usmansk county of Voronezh governance (Knyazhinskii, 1924, according to Ternovskii, 1929) in the present century. Along with this, the eyewitness account of A.P. Volynskii who wrote in 1736 and 1738 that wildcats of **a genus distinct from Astrakhan cats’’ existed in Voronezh governance (Kirikov, 1959) is wholly trustworthy. 309 310 414 conditions for the survival of wildcats in the territories under consideration, some 200 to 300 years ago, were far more favorable than at present. This is particularly true of the region between the Dnepr and Northern Donets and the Don basin. This was particularly so during the remote past (Middle Ages; Heptner, 1959). In any case the outline of the range in the Russian plains is quite typical of forms of western origin. References by some older zoologists to the occurrence (cited from Ognev, 1935) of cats in the former Belozersk county of Novgorod gover- nance, former Vytegorsk (Vishegorsk in Ognev, 1935 is a misprint) county of Olonets governance, former Shenkursk county of Arkhangel’sk governance, and finally to the southern (Zlatoustovsk) Urals are neither trustworthy nor confirmed. Intrusions at medieval Pskov (eleventh to fifteenth centuries; Tsalkin, 1952) pertain not to wild but obviously to domestic cats. In the last century the range shrank to insignificant remnants. This occur- red probably due to changes in environmental conditions and the general scarcity of game on which wildcats feed. It was not exposed to special direct pursuit. The chronology of this process is not clearly understood. Thus it has been suggested that all the data on wildcats of Latvia and probably also of Lithuania for the nineteenth century and possibly even the end of the eighteenth, pertain not to wildcats proper but to domestic cats turned feral (Paaver, 1965). In any event, cats disappeared earlier in Latvia than in Lithuania. In the stage of range contraction preceding the present one, 1.е., in the nineteenth century, at places even in the present century, the eastern boundary commenced evidently somewhere in Belorussia, in the Minsk or Mogilev region. To the south the easternmost points of habitation of forest cat extend- ed into the Kiev district to the west and southwest of Kiev (Radomyshl’, Stavishchi to the south of Belaya Tserkva), although it was evidently encountered right in Kiev. From these places the boundary turned in the direction of the Podol’sk district where the forest cat was recorded particularly in the region of Ushitsa in the basin of the upper Dnestr. From there the range boundary descended southward, encompassing the region between the Dnestr and Danube, to its eastern boundary at the swampy estuary of the Dnestr. It is possible that forest wildcats in the nineteenth century extended even farther east and lived, for example, in Chernyi forest near Kirovograd and in swamps and other places in the Dnepr valley, but sufficiently reliable data are not available. In the 1940’s to the 1960’s cats were reported from the western half of, and northern (Poles’e) Volyn district, the extreme north of L’vov, and from the northern part (Poles’e) of Roven and Zhitomir, and probably was encountered in the southern part of Zhitomir district and some places in ©0799 000099 Fig. 184. Range (reconstructed) of European forest wildcats, Felis (Felts) silvestris silvestris and F. (F.) s. caucasicus (Caucasus) in the Soviet Union. A—boundary of maximal reconstructed range (sixteenth to the nineteenth centuries); B—range boundary in the west in the 1940’s to the 1960’s (see Fig. 185 for details of the range in Moldavia). Question mark in the west pertains to possible occurrence in the middle of the twentieth century in Izmail, Zhitomir, and Kamenets-Podol’ districts and in Dnepr swamps. The following are some individual places of occurrence in the past: 1—‘*Kaluga governance’’; 2—‘‘Voronezh гоуетапсе”; 3— Kursk; 4— Putivl’; 5 — Sumy; 6— Aktyrka; 7 — Belgorod (together with No. 3, ‘‘Kur-Belogorod territory’); 8— Shipov forest at Pavlovsk on the Don; 9 —Zmiev; 10—Chernyi forest; 11 —Оперг swamps; 12— Mogilev. Crosses denote some erroneous references (V.G. Heptner). 416 Fig. 185. Distribution of European forest wildcat, Felis (Felis) 5. silvestris in Moldavia. 1 —Kagul swamp (lower Prut); 2—broad-leaved forest of Kodr; 3—northern Moldavian forest; 4—— floodplain forests and swamps of lower Dnestr (Yu.V. Averin and M.N. Lozan—data for 1967). Kamenets-Podol’sk. Wildcats were generally found in all the montane forests of the Carpathians within the Trans-Carpathian, Stanislav, Drogobych, and Chernovits districts, in the greater (southern) part of L’vov, and in the southeastern part of Ternopol’sk (Tatarinov, 1956). In Moldavia forest wildcats are distributed in forests along the Prut (almost to the mouth), at places along the Dnestr, and interfluve forests in the so-called Kodr (in Strashensk, Orgeevsk, Teleneshtsk, and Kotovsk regions; Kuznetsov, 1952; Yu.V. Averin and M.N. Lozan; Fig. 185). Evidently they also live in Dnestr swamps. There is no information relating to the far south of the interfluve of Dnestr and Danube (Izmail’ district); occurrence is generally shown, however, as ‘‘swamp forests’’ of the Dnestr and Danube (Sokur, 1960).*! ‘lThe map given by this author does not confirm these areas, referred to in the text as occurrences in the Dnepr estuary. 311 417 Forest wildcats have not been reported from Belorussia during the present century. Only one intrusion has been recorded for Minsk, in 1927. The animals are not even seen in Belovezh Forest where they were very common at some places. However, their appearance is still possible in southern Poles’e from adjoining sections of Volyn, Roven, and Zhitomir districts. In the Caucasus wildcats are distributed extensively in all forest regions of the Great and Little Caucasus. In the latter they occur in the east to Shusha meridian; they are absent only in the extremely high mountains, in the forestless mountains of Dagestan, and in the open steppes and semisteppes of the eastern Trans-Caucasus. In the Cis-Caucasian plains forest cats are present along the forested sections of the Kuban valley and encountered among reeds of the Kuban delta. They likewise live in forests (Paraboch) and shrubby regions along the Terek, at least to Kizlyar, but probably to the sea. The animal has been sighted in the Kuma valley down to its middle course, in the Sulak valley, and in the forests of Stavropol’. A reference to the presence of the forest cat in Talysh (Radde, 1886) has been questioned (Satunin, 1905; also Smirnov, 1922) since the skull brought from there proved to be that of domestic cat. Yet even now (Bobrin- skii et al., 1944 and 1965; and others) wildcats are nonetheless reported for the Talysh and Lenkoran. However, since the 1880’s no sightings other than the first report mentioned above are known from this part of the Caucasus and the animal was/is evidently absent there. Only the jungle cat is definitely known to occur in this region. The range of the forest wildcat in the Caucasus underwent no significant changes in the period under consideration. Thus, in spite of a reduction in forests in the Stavropol’ upland, cats still lived in Temnolessk forest 40 km south of Stavropol’. Only localized reductions occurred due to deforestation. However, this species is also found in gardens and vineyards with sparse forests and shrubby thickets (Kizlyar for example).*? Geographic Range outside the Soviet Union The range outside the Soviet Union (see Fig. 181) encompasses Europe in the west, including England (absent in Ireland and evidently never present “Range according to Bikhner, 1905; Satunin, 1905 and 1915; Dinnik, 1910; Brauner, 1914, 1923, and 1928; Migulin, 1928; Charlemagne, 1928; Ternovskii, 1929; Fedyushin, 1930; Ognev, 1935; Heptner and Formozov, 1941; Vereshchagin, 1947 and 1958; Kalnin’sh, 1950; Heptner, Turova and Tsalkin, 1950; Kirikov, 1952, 1959, and 1966; Korneev, 1952; Kuznetsov, 1952; Serzhanin, 1955 and 1961; Tatarinov, 1956; Sokur, 1960 and 1961; Lozan and Korchmar’, 1965; Paaver, 1965; Alekperov, 1966; Korchmar’, 1968; and also original data of Yu.V. Averin, М.М. Lozan, V.G. Heptner; and others. Biz 418 there), northward to the Baltic Sea, and southward to southern Spain, Italy, Sicily, the Balkans, and all of Asia Minor. These cats are not known to occur in Syria and Iraq but may perhaps intrude into areas adjacent to Turkey. They have not been reported for Iran and are evidently absent there. Cats inhabiting islands of the Mediterranean Sea (Corsica, Sardinia, Crete, and the Balearic Islands), sometimes even those of Italy and Sicily, are placed in this group, and/or libyca group, or considered a transitional group. Finds of Pleistocene cats assigned to this species generally do not occur beyond the limits of the present-day range. Wildcats have been found in Holocene Crimea (Birulya, 1930)* and in cultural horizons of some five thousand years ago in southern Scandinavia (Scania; Eckman, 1922)*, and probably extended to the Gulf of Finland in the Pri-Baltic during the Holocene (Paaver, 1965)*. The depiction of the Holocene (*‘post-glacial’’) range as having encompassed the eastern part of the Don basin and central Volga, including also Trans-Volga (Haltenorth, 1953) is evidently based on assumptions. There is no concrete proof of this range extension in our literature. : In all probability forest wildcats were not distributed beyond the Don basin and perhaps not even its western part. In any case cat remains from Sarkel (lower Don, ninth to the twelfth centuries) have been adjudged those of domestic animals (Vereshchagin, 1959). Evidently European and Caucasian portions of the range may be regarded as not contiguous across the Russian plains. This is indicated by the independent status of the subspecies of the Caucasian form and its affinity to animals from Asia Minor. Forest wildcats intruded into the Caucasus from Europe, where they were autochthonous (Kurten, 1965), through the Balkans and Asia Minor in the Pleistocene. (V.H.) Geographic Variation Several subspecies have been described for the forest wildcats of the group defined here; however, they have not been satisfactorily characterized to date, and have been separated for the part without adequate basis. Although geographic variation within the Soviet Union still requires clarification, two forms may be tentatively recognized in the USSR. 1. Central European forest wildcat, Е. (F.) 5. silvestris Schreb., 1777 (syn. ferus, ferox, and euxina). General color quite dark with a gray tone; pattern on head, dorsal spinal band, and transverse stripes and spots on trunk distinct and usually vivid. Rings on tail may range from three to ten, but more often seven in number. *Not in Literature Cited— Sci. Ed. 419 Body length (20 animals; western Ukraine, Carpathians) of males 480-910 mm (М 633) and of females 400-690 mm (М 540); tail length in males 330-350 mm (M 313) and in females 280-340 mm (M 294) (Suminskii, 1962).13 Data on the measurements of forest wildcats from Moldavia and adjoin- ing places are scanty and somewhat contradictory. The average body length of three males from Moldavia has been reported as 810 mm and of four females as 750 mm; tail length in males 360 mm and in females 355 mm; length of hind foot in males 160 mm and in females 124 mm; height of ears in males 60 mm and in females 49 mm; and weight of males up to 8.0 kg and of females to 6.5 kg. Greatest length of skull in males (three) 87.8-91.8 mm; condylobasal length of skull in males (five) 81.2-104 mm and in females* 80-91 mm; and zygomatic width in males (six) 62—78 mm and in females* 61-90 mm (Yu.V. Averin and M.N. Lozan; Mindlin, 1938)**. According to other data (Brauner, 1928) animals from former Bes- sarabsk governance, former Odessa county (one specimen from Podolia) had the following measurements (males and females together): body length (eight) 520-630 mm (M 590); tail length (eight) 310-340 mm (M 320); length of hind foot (five) 125-140 mm (М 133); and height of ears (five) 60-63 mm (M 61). Condylobasal length of skull (six) 83-113 mm (M 98.7); muzzle width above canines 22—31 mm (М 26.6); interorbital width 31.0-43.2 mm (М 36.4); zygomatic width 64.0-86.5 mm (М 75.1); and length of upper tooth row 21-30 mm (М 26). ` Greatest length of skull (mean, Poland, 10 males and females) 97.4 mm; condylobasal length 88.3 mm; zygomatic width 69.1 mm; interorbital width 19.8 mm; and postorbital width 34.9 mm (Suminskii, 1962). Greatest length of skull of fully adult animals from central Europe (Alsace and western regions of former German territory): males and females (25) 88-110 mm (M 94.1); greatest length of skull in males (6) 96-110 mm (M 102); condylobasal length in males (26) 86-101 mm (M 91.6) and in females* 80-89 mm (М 84.7); and zygomatic width in males (26) 66-80 mm (M 71.9) and females* 63-69 mm (M 65.3). Weight: males and females (189) 3,750 to 11,500 g (M 6,880); males (108) 5,000 to 11,500 g (M 7,180) and females (33) 3,750 to 10,000 g ‘2Information on this cat is almost nonexistent in Soviet museums and data in Soviet literature scanty. Information derived from foreign literature and from adjoining or nearby territories has been used to characterize it. Data by Suminskii (1962) on weight are extremely exaggerated (males 5.9 to 14.8 kg, etc.) and have not been included here. *Number of females omitted in Russian original—General Editor. **Not in Literature Cited —Sci. Ed. 313 420 (М 5,970) (Haltenorth, 1957). Although these figures do not include extreme values (see above), they are nevertheless sometimes considered, not without reason, to be on the high side. High values are usually attributed to hunters’ errors. Thus the weight of 36 pure-blooded adults, in part extremely large animals from Slovakia, weighed and measured by a zoologist (Sladek, 1966) was: males (20) 3,300 to 7,700 g (M 5,248) and females (16) 2,600 to 5,840 g (M 4,233). The largest male in this series had a body length of 880 mm, tail length 310 mm, length of hind foot 145 mm, and height at shoulders 430 mm. Greatest length of skull was 107 mm, condylobasal length 96 mm, and zygomatic width 76 mm. Found in the extreme west of the European part of the Soviet Union. Outside the Soviet Union this form is encountered in the European part of the range of the species, except in Scotland and islands of the Mediter- ranean Sea. 2. Caucasian forest wildcat, F. (Е.) 5. caucasica Satunin, 1905 (syn. daemon and trapezia). General shade of color gray, on the average somewhat lighter than in nominal form; pattern on head and dorsal band well developed, but pattern on trunk and partly on tail reduced. Transverse bands and spots on trunk usually faint or absent, but well defined in a few individuals. In addition to black tip on tail, only three distinct black transverse rings usually present. ‘‘Pelage rich, dense, and soft. On the back an average of 8,900 hairs occur in a 1 cm? area, and 2,000 in the same on the abdomen; ratio of underfur hair to top hair, 10 : 1. Guard hair stand somewhat above top hair. Latter comparatively short and do not rise much above underfur. Average length of guard and four categories of top hair on the back: 44, 41, 39, 37, and 36 mm, and thickness 81, 87, 71, 62, 43, and 20 microns; underfur hair 34 mm and 20 microns respectively. Hair length on abdomen less than on back’’ (В.Е. Tserevitinov).4 Body length of the adult cats usually 70 to 75 cm, height at shoulders 26 to 28 cm, and height at sacrum 30 to 32 cm. Weight often 5,200 to 6,000 g and up to 8.0 kg; rarely even more (Dinnik, 1914; Satunin, 1915). Two adult males from Caucasian preserve weighed 5,610 to 6,200 g and three females between 4,150 and 5,000 g. Found in the Caucasus. Outside the Soviet Union, found in Asia Minor. Melanistic individuals are encountered among Caucasian cats and have been separated into a distinct species, F. daemon. Actually these are simply 44 Тре information given here differs significantly from data given above for central European cats. This discrepancy is evidently due more to differences in methodology than to differences between the two subspecies. The data of B.F. Tserevitinov in any case are wholly comparable to that given for other Soviet cats. 314 421 feral domestic cats (material from the Zoological Museum, Moscow Uni- versity) or, perhaps, hybrids of wild and domestic cats. Occasional cases could well be melanistic wildcats, since some geographic localization of these forms is seen in the eastern part of the southern slope of the Great Caucasus in the districts of Lagodekh, Zakatal, and Nukhi (former Nukhinsk county and former Zakatal’sk area). Data are available for the last century about the presence of such forms in the eastern part of the Little Caucasus from Kirovabad (former Elisavetpol’) to the upper Terter and Shusha (Satunin, 1915). In the middle of the present century they have generally been caught in the same places in Little Caucasus, namely, Stepanakertsk and Shushinsk regions of Azerbaidzhan (Karabakh range, two to four animals in winter; Alekperov, 1966). In 1948 in a fur warehouse in Baku, of the 117 skins of forest wildcats 14 were pure melanistic and 3 dark brown (A.P. Korneev). Evidently these figures represent all of Azerbaidzhan. The Caucasian wildcat is a distinct well-characterized form and its range readily demarcated from that of western forest forms. Unfortunately a comprehensive taxonomic analysis has not been attempted to date. However, it has always been thought that this form is larger than the European form, which is not correct. The main difference between the two lies in the color characteristics described above. One of the features of the Caucasian form, compared with the western, is that no hybridization of the former with domestic cat takes place, which is a very Common occurrence in the western form. There is no information from the Caucasus about extensive hybridization, let alone hybrid popu- lations. Living conditions are not favorable for hybridization. Extensive development of suitable landscapes and extensive forests (Kuban Caucasus, and a series of other areas) ensure a natural population in numbers and composition. Figures relating to the admixture of blood of domestic cat in the Caucasus (see above) are no doubt erroneous and reflect imperfect methodology. Outside the Soviet Union, apart from the two forms described above, the following subspecies of European forest wildcats are usually recognized: 1) F. (F.) 5. grampia Miller, 1907—Scotland; 2) Е. (F.) 5. tartessia Miller, 1907—southern Spain; 3) F. (F.) 5. molisana Altob., 1921 —Иау; 4) Е. (Е.) 5. euxina Pocock, 1943—Dobruja; and 5) F. (F.) 5. тогеа Trouess., 1904—-southern Greece (Ellerman and Morrison-Scott, 1951). Only the subspecies grampia would appear genuine (Haltenorth, 1957). Thus there are only three subspecies of forest wildcats. The view that 422 (€961) AoquAY рие AOJOY рие “(ээцэтб Jo AWapeoy ‘шпэзпи| [е180|007, wo $1эщ1994$ — CEG) лэи8 О ‘(CT6T) ащес ‘(Фтбт) уши jo вер ‘АтзлэлиГ] м09$0И] ‘шпэзпИ| [29130007 Шол] $1эщ12э46 :9}0N Gaal ост 96 TZ 0`05 See ЭС РС See 9 0`05 | ба 681 Cale 0O'9oT 06 0:99 $ 0`09 TZ [198 0`00Т 708 (LG Стб 9901 €L8 €? cer СРТ ОЕТ $ 505 05$ OLT Ё [65 059 OVS L W хеш UTUL и зэтешеэН хеш зе JOT 50$ Ele SOT aa) 0°88 ‘96 ОРТ 06 059 ии Ц1001 [ет5звилео Joddn jo ц}3иэ] MOI 100} 1э44п до ц}8иэ] YIPIM [еифо0}504 YIPIM |еи9золэи] рим эцешоЗА7 41849] jeseqojApuodg [[NYs JO yBua] 1$3]еэ19) 100} ри дю yysuaT yysuay пет, Ч184э| Apog $эхэрц] 1221502102 ‘5 (J) “YY Чеэрим 35эл0у ULISEINeD Jo [NYS pue Ароа Jo $зиэшели$ез | “py 7qe 1 elt 423 there can be no geographic variation due to hybridization of forest wildcat with domestic cat is not correct (Suminskii, 1962). The forms listed above are typical for the group si/vestris. The following forms occupy an intermediate (transitional) position in relation to the /ibyca group: 4) F. (F.) 5. jordansi Schwarz, 1930—Balearic Islands; 5) Е. (F.) 5. reyi Lavaud, 1929—Corsica; 6) Е. (F.) 5. cretensis Halt., 1953—Crete; and 7) F. (F.) 5. sarda Lat., 1885—Sardinia, Sicily, northern parts of the North Africa from Cyrenaica to Morocco and southern Atlas, and Algerian Sahara (Haltenorth, 1953 and 1957). Often, all of these forms are combined into one (sarda) and placed in the libyca group (species). (V.H.) Biology Population. In the western parts of the range, the forest wildcat became scarce even in the early nineteenth century in Latvia and almost extinct in Lithuania and much of Belorussia. They are, at present, extremely rare and only single individuals are caught in Lithuania and Belorussia (Poles’e). About 1,000 cats still live in the Carpathians and over a hundred are Fig. 186. Montane forest in the Carpathians. Habitat of forest wildcat and lynx. Vicinity of town of Rakhovo. August, 1967. Photograph by G.I. Simkin. 315 424 caught annually in the Trans-Carpathians and the western part of Volyna; cats are now extinct elsewhere in the Ukraine. In Moldavia forest wildcats are common in the lower reaches of the Dnestr and Prut but rare in the Kodr. In the early 1960’s from 20 to 40 cats used to be caught every season in this region. Forest wildcats are quite common over large areas in the Trans-Caucasus and the Caucasus, in the north to deltas of the Kuban and Terek. They are particularly numerous along the Black Sea coast but rare near the Caspian Fig. 187. Ancient beech forest. Habitat of forest wildcat in Babich area. Carpathians. Photograph by Yu.D. Tret’yak. ЛЯ 425 [Sea]. In Azerbaidzhan these animals are caught in the thousands, in Armenia, Georgia, and Dagestan in the hundreds, and in the Groznen district and Krasnodar territory in the northern Caucasus again in the thousands (up to 5,000). In the forested ‘“Chernye Gory’’ range—the first ridge of the mountains —wildcats were numerous in the central part of the northern slope of the Great Caucasus in the 1920’s (V.G. Heptner). Habitat. In the western regions of the Ukraine wildcats inhabit broad- leaved hornbeam-oak, beech, and mixed forests on the Volyno-Podol’sk plateau and in the Carpathians. They are absent in the spruce taiga along the crest dividing the Carpathians, and the subalpine belt of these mountains. Along particular slopes of the mountains they ascend to the upper boundary of beech and mixed forests at a height of 1,200 to 1,400 m above sea level. The main habitats are beech forests on the southwestern slopes of the Carpathians, i.e., the old “‘preserved’’ forests growing at the upper boundary of wooded vegetation. These forests consist of massive, centuries-old beech and sycamore trees, which yield an abundant crop of seeds every two or three years. Many trees have hollows in which not only birds but also squirrels, martens, and cats nest. In these forests mouse-like rodents and birds are abundant; wildcats mainly hunt the first of these (F.I. Strautman; Konyukhovich, 1953; Taratinov, 1956a). In Moldavia wildcats inhabit beech forests as well as swamps consisting of dense thickets of reeds and tala willow with some hollow willows and black poplars growing among them. Such swamps abound in small tributaries and lakes. In the swamps of the lower Prut there are no forests in the floodplains but wildcats are found there in large numbers. They live in the hollows of old willows along dams and on ridges, in reed thickets along banks of reservoirs, and on floating islands (Lozan and Korchmar’, 1965). The habitats of these cats are particularly diverse in the Caucasus. They are commonly found on ridges, islands, and in dense reeds in the swamps of the Kuban and Terek, and quite often encountered on islands that have broken off and are floating far from shore. There it leads a life similar to that of jungle cat. In Dagestan forest wildcats are encountered in reeds along the sea coast (Dinnik, 1914) and in Georgia regularly visit marine surf zone (M.P. Pavlov). Along the southern slope of the Great Caucasian range, they live down to the very shore of the Black Sea where they are confined to thickets of cherry laurel, yews, and box trees (A.A. Nasimovich). Wildcats are common, at places even abundant, along montane slopes overgrown with beech and oak. In the Caucasus they prefer the broad-leaved forest zone, especially sections which adjoin the glades and valleys of rivers, and inhabit mainly the central montane belt. The cat is met with significantly more rarely in dark coniferous forests, and in the subalpine belt it occurs only as an exceptionally rare occurrence. Thus in Caucasian preserve 76.7% 316 426 Fig. 188. Old beech forest with vines. Habitat of forest wildcat on Black Sea coast of the Caucasus. Photograph by A.P. Zhandarmov. of occurrences took place in the belt of broad-leaved forests, 13.6% in dark coniferous forests, and 9.7% in the subalpine belt. Instances of occupation of the deep montane regions covered with coniferous forests are few; the cat is confined exclusively to river valleys usually that support broad-leaved species (Teplov, 1938). The above distributional features of cats according to habitat depend on availability of food, primarily its accessibility during the winter period. In montane forests cats show preference for cliffs or rocky sections. They invariably prefer rock cliffs to hollows of trees and set up their dens 318 427 far more rarely in the latter than among rocks. In the forest, except for rocky sections, they select impenetrable thickets and places carpeted with fallen trees and branches. In shrubs and sparse forests they usually select places with outcrops of rock slides, stones, and precipices (Dinnik, 1914). In years of mass breeding of rodents in the steppes, cats sometimes descend the foothills to catch voles in fields (see below). At such places, inhabited also by jungle cat, wildcats live in the ‘‘upper’’ forests along the slopes and not in the valleys (Zakatalo-Nukhinsk valley; N.N. Rukovskii). Most of the population lives in the mountains at a height of 300 to 600 m above sea level and even higher (600 to 800 m) at some places; when intruding into the subalpine belt, however, they sometimes ascend up to 1,500 to 1,800 m above sea level (Dinnik, 1914). In Armenia wildcats live in the forest regions at a height of 700 to 2,500 m above sea level (Dal’, 1954). In some parts of the Caucasus this cat lives in extremely wet places, apart from river mouths (see above). For example, in the Kolkhid lowland (Batumi and Poti) rains are very frequent, with downpours almost every day in autumn and winter. Local cats are not afraid of dampness and wander day after day in mud, wet grass, or weeds, which drip water on them. They also enter and swim through rivulets and brooks. During high floods, when tens of kilometers of lowland forests and swamps are inundated, they live BT, Fig. 189. Beech forest on the southern slope of Great Caucasus range. Biotope of forest wildcat. Azerbaidzhan. November, 1957. Photograph by N.N. Rukovskii. 318 428 in trees for weeks, feeding on Norway rats taking refuge there (М.К. Vereshchagin). A wildcat will not avoid the proximity of humans and often lives not only close to settlements but even in granaries, barns, attics, summer houses, and larger residences. The European forest wildcat is not adapted to living in regions with deep, loose snow cover. With a high weight load on the resting surface of the foot at 88 to 118 g per cm/?, it flounders and advances with great difficulty on loose snow since it is relatively short-legged (height of forelegs and lower surface of chest to the ground in males 25 cm, and in females 22 cm). Moreover, it cannot get at rodents under the snow. These are evidently the reasons why the cat does not ascend high into the mountains, especially in winter. In Caucasian preserve with a deep snow cover the cat descends the mountain slopes, avoiding regions of deeper, more abundant snow depth which are difficult to negotiate and where birds are scarce (A.A. Nasimovich). The wildcat usually inhabits only regions where the mean of winter snow cover does not exceed 10 to 20 cm (Formozov, 1946). This could explain its absence in the central zone of European Russia (depth of snow cover 40 to 60 cm), not to mention more northerly regions. Fig. 190. Thickets of sea buckthorn, with common buckthorn, blackthorn, and dog rose. Habitat of forest wildcat. *‘Chernye Согу”” south of Ordzhonikidze, near Balta. May, 1966. Photograph by Т.Р. Mitina. 320 429 In unusually severe, abundantly snowy winters forest wildcats suffer starvation and sometimes even die of emaciation. For example, in the winter of 1953/1954, in the Kuban and in all of the southern European part of the USSR, there was unusually severe, prolonged and constant subzero temperatures together with excessive snowfall. The minimum air temperature was —21° С in November, —24° С in December, —26° С in January, and —35° С in February. Snow drifts in swamps were 2.0 т deep and at places even more. This winter proved fatal to many inhabitants of these swamps. In February, exhausted and emaciated wildcats, foxes, and wild boar were observed to abandon their usual caution and moved onto farms during the day; cats often attempted to hide in cattle sheds. Not only a large number of small animals and birds died, but predators also suffered from hunger and died of emaciation. In March, 1954 one wildcat, three foxes, and two minks [Mustela lutreola] were found dead (Pavlov and Kiris, 1956). Wildcats often endure starvation and die of emaciation during winter even in Central Europe (Haltenorth, 1957). In the very severe winter of 1962/1963 a deep snow cover deprived wildcats, especially those living in montane regions, of mobility, crippling their search for rodents. They descended into farmsteads. In February, 1963, dead cats with empty. stomachs were found (Miiller-Using, 1963). Domestic cats are capable of surviving total starvation for 19 days. Food. \n the Carpathians wildcats mainly feed on the yellow-throated mouse [Apodemus flavicolis], red-backed vole [Clethrionomys rutilus| and ground vole [Pitymus subterraneus| which occur in large numbers during *“‘mouse’’ years in beech forests (F.I. Strautman); they also catch field hares [L. europaeus] (Tatarinov, 1956). In the Trans-Carpathians they feed mainly on mouse-like rodents, galliform birds, and squirrels (Konyukhovich, 1953). In the Dnestr swamps they hunt for small voles [Microtus], water voles [Arvicola], and birds (Brauner, 1923); in the Prut swamps, judging from an analysis of 77 food samples, they feed mainly on water voles (91% of samples, Norway rats (24%), and muskrats (13%). Of the birds available, they most often attack warblers (52%), white-eyed pochards [A ythya пугоса] (20.8%), coots [Fulica ата] (9.1%), spotted crakes [Porzana porzana]| (5.2%), and gadwall ducks [Anas strepera] (2.6%). Moreover, they eat wood mice [Apodemus sylvaticus] (3.9%), fish (3.0%), mollusks (2.6%), and beetles (1.5%). In Moldavia, in the lower courses of the rivers, rodents predominate in the winter diet while birds, fish, and crayfish predominate in the summer diet. Wildcats catch a very large number of aquatic and other birds in May and June when most of these birds are sitting on their nests (Lozan and Korchmar’, 1965). In the Kuban delta the main food of wildcats consists of Norway rats and water voles, which are abundant there, and additionally muskrats and waterfowl (M.P. Pavlov and L.A. Volyanskii). In the northern Caucasus the 430 319 Fig. 191. Thickets of willow, blackberry, blackthorn, and reeds. Biotope of forest wild- cat in Terek valley, northwest of Groznyi. May, 1966. Photograph by I.P. Mitina. main prey of this cat in summer are mouse-like rodents and edible dormice [Glis 215], and more rarely birds. From time to time various fruits, including mountain ash, have been found in stomachs. On rare occasions it will attack young roe deer and chamois (Dinnik, 1914). In Dagestan broods of pheasants and partridges suffer most from this predator (Heptner and Formozov, 1941). On the Black Sea coast, judging from an analysis of food remains, it feeds on small birds, mice, shrews, and hares. On one occasion the feathers of a white-tailed eagle [Haliaetus albicilla|, and on another the skull of a kid, were found in dens. When necessary, it eats a fresh kill but usually approaches it only after jackals have begun to devour it. With the appearance of a wildcat, jackals withdraw from the carcass and return to it only after the cat has left (Obraztsov, 1928). In Caucasian preserve (12 stomach contents and 35 feces, as percentage of total number of both) the food of wildcats is as follows: mammals 100, rodents 95.7, mouse-like rodents 93.6, bush voles [Pitymys] 76.6, wood mouse 14.9, edible dormouse 2.1, European hare 2.1, mole [Talpa] 4.2, shrew 4.2, chamois 2.1, small birds 3.5, insects 2.1, and sedge and grass (leaves) 431 10.6 (Teplov, 1936). Chamois remains were found in feces collected on December 22; the cat evidently ate carrion. Similarly, remains of a drake mallard eaten by a cat were also seen. The stomachs of three cats caught in August and September in Zakatalo-Nukhinsk valley contained yellow- throated mice and edible dormice; the remains of a rock partridge were found near a den. In the stomachs of cats caught in summer in the Ismaillinsk region of Azerbaidzhan only the remains of yellow-throated mice, wood mice, and edible dormice were found (Rukovskii, 1953). During the drought of 1948 many wildcats and jackals were sighted on the Black Sea coast around drying lakes (for example, Lake Imanati, upper Gurinki River), which supported a large number of nutrias and aquatic birds. These predators regularly hunted for rodents and birds. In January, 1949, after the freezing of pools unaffected by drought, the cats turned to nutrias hiding in bushes in large numbers as they were compelled to live on land. In 1939 in Abkhaz’ya some 60 wildcats and 70 jackals were extirpated during the summer over a comparatively small area of the Inkit swamp. It is not surprising that with such a huge population of predators in Georgia after the severe winters of 1948/1949 and 1949/1950 nutrias everywhere became rare. In the stomachs of four cats caught in December, 1949, near Lake Inkit there were remains of mallards, sheldrakes | Tadorna tadorna], song birds, water voles, and small mouselike rodents (М.Р. Pavlov). Cats caught on June 12, 1937, in Armenia in a beech forest had filled their stomach with lizards (E.P. Spangenberg). In western Europe the cats feed on rodents (hamsters, Norway rats, dormice, water voles, voles, and.wood mice) and from time to time on small predators such as martens, polecats, ermines, and weasels in addition to shrews and moles. They do not relish insectivores. The remains of squirrels and forest birds constituted an insignificant quantity in stomach analyses (Satunin, 1915; Beaufort, 1961; Traubot, 1961). Wildcats there also attack, from time to time, the fawns of [red] deer, roe deer, and chamois. They hunt hares and rabbits and catch fish, inflicting some damage during spawning, especially on trout (Gaake, 1901*; Dinnik, 1914; Haltenorth, 1957; Beaufort, 1961). Individual cats specializing in catching the young of ungulates are also encountered. When living close to human settlements, often right inside them, this predator can cause significant damage to poultry. From the foregoing data on the food of the forest wildcat in different parts of its range it may be concluded that small rodents (mice, voles, and rats) constitute their primary food everywhere; birds (chickens, ducks, and more rarely passerines) occupy second place, followed by dormice, hare, *Not in Literature Cited— Sci. Ed. 432 nutria, muskrat, and insectivores. The predator hunts other animals in years of low population of mouse-like rodents, or incidentally. While hunting, the cat wanders throughout the forest and along the edges of forests and glades. Where conditions are favorable, the forest wildcat will readily feed in a field. In the winter of 1923-1924 the weather was free of snow in December and January around Vladikavdaz. It was a year of high numbers of small rodents, mainly gray voles [Microtus]. Cats, also abundant that year, left the forests in the foothills (Chernye Gory), together with foxes, during twilight hours and entered singly into fields adjoining the forest. Even wolves came. A group of hunters shot several wandering cats and foxes in the brief twilight period. The cats were very obese (V.G. Heptner). In pursuing prey, sometimes the cat climbs to the very top of high trees, even jumping from one tree to another. It lies in wait for a prey and catches it by executing a few leaps. Sometimes it lies in wait standing (lying) near a burrow or crevice among rocks. Its leaps can reach a span of 3.0 m. The cat relies on vision and audition in searching for prey. Olfaction is relatively weakly developed. It hunts for nutria and waterfowl from trees overhanging the water in which they swim. These cats have been observed hunting in Georgia during the dawn hours (Pavlov, 1953). On one drying lake a cat attempted to attack an adult пила feeding in a strongly shoaling and boggy section of the shore. The cat approached the nutria by executing large bounds but soon mired in the mud and had to return to its former place of ambush (М.Р. Pavlov). If the prey is small, the cat grabs it in its claws and then kills it by piercing the neck or occiput. There are references to this cat jumping onto the back of large animals and attempting to bite the neck or carotid (Bikhner, 1905; Haltenorth, 1957). Failing to capture a prey in its initial attack, the cat does not continue the attempt. In the stomachs of cats caught while hunting, usually three to seven mice or voles have been found, but in ‘‘mouse’’ years significantly more. In Caucasian preserve on March 8, 1934, the stomach of one cat contained 16 rodents and one tit [Parus]; their total weight was about 270 g. In the stomach of another cat 26 mice were found which weighed about 500 g (Satunin, 1915). — Captive wild kittens aged 1.5 to 2.0 months are daily fed up to 10 mice each (about 180 g) while year-old animals receive an average of 900 g of horse meat every day (Teplov, 1938). Home range. Not much is known about the dimensions of the individual home range of a wild cat. In the Prut swamps a cat or a whole litter lived on every third or fourth floating island, comprising an area of 1.0 to 2.0 hectares (Lozan and Korchmar’, 1965). In Caucasian preserve, in the proximity of the Kishinsk cordon, no less than 20 animals, or an average of one cat per 60 hectares, were counted in the autumn—winter season of 1936/1937 in a 322 433 total area of about 12 km? (Teplov, 1938); in the Caucasus there are many regions where the population density of this predator is significantly higher. While living in a given section the cat leaves scent marks at different sites. The number of such marked sites greatly increases during the period of estrus when the preanal glands of males and females enlarge and liberally secrete a compound containing trimethylamine and other odoriferous sub- stances. During the rut the male moves around trees with its tail raised and sprays its scent on the bark. Some cats begin to mark their territory when 160 to 170 days old; the majority commence territorial claims only after 365 days of age (Haltenorth, 1957). Burrows and shelters. In the Carpathians and Trans-Carpathians wild- cats live in the hollows of beech trees and in rocky fissures (F.I. Strautman; Konyukhovich, 1953). In the swamps of the Dnestr and Prut they live in summer in forks in the trunks of old willow trees, in abandoned heron nests, and in the hollows of willow trees and black poplars. In the Kochul’ swamps (Moldavia) most cats live on floating islets. Usually the den with kittens is located in the central part of the island among densely grown reeds. Well-beaten pathways lead to it. Beside the den can be seen areas in which the kittens play and where prey is consumed (Brauner, 1914; Lozan and Korchmar’, 1965). In the Caucasus, depending on the locality, these cats sometimes prefer to live in crevices among rocks, sometimes among hollows in trees, Or sometimes in abandoned burrows of foxes and badgers (Dinnik, 1914). In Caucasian preserve the most common shelters are hollows in the trunks of fallen trees, and more rarely hollows of growing trees, rock crevices and caves, and abandoned burrows of badgers and foxes. On May 11, 1936 in a preserve near the Kishi River the den of a cat was found on the bank of a small creek in a thicket of alder, nut trees, and wild apple. It was located in the hollow trunk of a fallen rotten elm about 45 cm in diameter. The hollow was about 30 cm in diameter and 7.0 m long. There was no bedding except the powdery wood rot. In the nest were the kittens (Teplov, 1938). In Zakatalo-Nukhinsk valley wildcats live almost exclusively in the abandoned burrows of badgers, which are numerous on slopes covered with beech forest (2 or 3 burrows per 10 hectares). In this same valley, at the upper forest boundary where burrows are few, cats use rock fissures for shelter. Only once was a den found in the hollow of a fallen poplar. It is characteristic that in places where the cat lives mainly in burrows, it does not climb trees to save itself from dogs but attempts to retreat into its burrow. On several occasions even when dogs chased a cat and began to attack it, the latter did not attempt to rescue itself by climbing a nearby tree (N.N. Rukovskii). In taking up residence in the hollow of a living tree, the cat selects one 434 which is not located very high above the ground. A hollow open above is not selected. On one occasion a wildcat’s den was found in the center of a large firewood pile. The nest was made of dry grass and reeds and there were six very healthy kittens with opened eyes (end of May, 1884; Rossikov, 1887). It is evident that this cat does not dig a hole for itself. If the shelter is in a burrow or rocks, it is lined with dry grass, leaves, or bird feathers; while living in a tree hollow, the cat is satisfied with the wood dust present in it. It uses hollows or breaks as temporary shelters, or lies among dense vegetation. In years when the flea population is high in summer the cat does not live for long in the same den, but shifts from one to another. In winter, when soft and loose snow does not permit long-distant wandering, it remains for long times in the same den (Dinnik, 1914). In Caucasian preserve it uses the same site as a day rest for quite some time. In western Europe it selects hollows, rock crevices, and abandoned burrows of foxes and badgers for dens but sometimes makes an open nest in dense vegetation in marshes (Haltenorth, 1953). Daily activity and behavior. The wildcat mostly leads a crepuscular and nocturnal life. It usually sets out to hunt an hour or two before sunset and remains active throughout the evening and the first half of night, then rests, and reappears again only at dawn. In summer, however, it also hunts during the day. The stomachs of cats caught two hours before sunset contained freshly caught rodents. Many cases are known, when cats have stolen domestic fowl during the day and have been encountered by hunters. The wildcat is less cautious and fearless than other animals. It will permit a man on horseback to approach within 30 to 40 paces, after which it runs away unhurriedly. It swims well but avoids water when avoiding danger. A trapped cat rumbles and hisses at an approaching man but does not attack. A captive animal is very malicious and does not become accustomed to man; taming kittens is difficult indeed. According to observations made in the Caucasus, it invariably lives and hunts alone. According to data of earlier zoologists in western Europe, however, both parents live with the litter. Litters of young with two adult animals have reportedly been seen there. This information appears erroneous. The role of the male in raising kittens is still not known (Haltenorth, 1957). Seasonal migrations and transgressions. These aspects are not known. In Caucasian preserve the wildcat does not, as a rule, wander even though food may be scarce (Teplov, 1938). In the Carpathians population changes are partly due to cats moving on during years of “‘роог harvest’’ of mice (Е. Strautman). In western Europe cats descend from the mountains to the plains in winters of abundant snow (Miller-Using, 1963). Reproduction. In the Trans-Carpathians estrus takes place from January to March (Konyukhovich, 1953); on the lower Prut (Moldavia) in early March (Lozan and Korchmar’, 1965); in the northern Caucasus in February 435 (old calendar; Dinnik, 1914); and in Azerbaidzhan in the middle of February (Vereshchagin, 1942). However, according to recent observations based on captured pregnant females and the appearance of kittens, wildcats come into heat from December to February and May to July, and possibly in other seasons in different regions of the Caucasus. In Central, Europe animals in estrus are seen from the second half of February to the first half of March. Data are available which reveal a second breeding season, in autumn (Trauboth, 1961). In Scotland, between the end of May to early June, females often come into estrus a second time, and from September to October for a third time (Matthews, 1941). Females during repeated periods of estrus have enlarged corpora lutea from the first pregnancy, pointing to the possibility of two or three litters in a single year. Subsequent periods of estrus are irregular. It is not known whether the mother suckles the kittens of the second, let alone the third litter, and whether this is even possible. Quite likely a subsequent litter appears due to the death of the previous litter. In some females estrus continues for one to five days. Ovulation is induced [by copulation]. Spermatogenesis proceeds year-round (Haltenorth, 1957). Mature sperm were found in the vas deferens of a cat caught on November 12, 1957, in the Ismaillinsk region of Azerbaidzhan Soviet Socia- list Republic (N.N. Rukovskii). During estrus the preanal glands become enlarged in both sexes (see above) and sometimes vocalization, resembling that of domestic cats, is heard; both sexes are vocal. Fierce fights ensue between males during the rut. Gestation requires 63 days, sometimes up to 68 days (55 to 68 days in domestic cats). In Western Europe kittens of the first litter appear from the end of April to May. The second litter is seen at the end of July to early August, and the third (in Scotland) at the end of November to early December, and also mid-January (Matthews, 1941; Haltenorth, 1957). In the Trans-Carpathians kittens are born in April and May (Konyukhovich, 1953). In the Carpathians litters of four kittens with opened eyes were found in mid-May (F.I. Strautman; Tatarinov, 1956a); a juvenile cat with a body length of 30-40 cm was caught on November 15, 1952 (Tatarinov, 1956). In Moldavia, on the lower Prut, kittens are seen in the first half of May. On July 10 a cat was still suckling three grown-up kittens. On July 19 a young female had a body length of 63 cm, while a young male caught on July 25 weighed 1,900 g (Lozan and Korchmar’, 1965). In the northern Caucasus kittens are seen toward the end of March or in April (old calendar; Dinnik, 1914) and in Azerbaidzhan in April (new calendar). Kittens with unopened and opened eyes were found at the end 324 436 of May on the Malka River (Rossikov, 1887), and four kittens with eyes fully opened and milk teeth well developed were found on May 11, 1936, in Caucasian preserve. A female caught in Azerbaidzhan on February 23, 1953, contained five large fur-covered fetuses (three females and two males). The fetuses averaged 117 mm in size and the weight of the females was 68.6 g and that of the males 72.0 g. As these kittens would have been born toward the end of February, this cat must have mated in the second half of December. A female caught on February 21, 1953 in the same region had four implantation sites 43 mm in diameter and fetuses weighing 332 mg. It evidently mated in mid-January and the kittens would have been born in mid-March. Lactating cats were caught on June 12, 1937 at Sevan (Armenia) and on June 21, 1939 in the Kubinsk region of Azerbaidzhan. A female caught on May 16, 1951 in the Zakatalo-Nukhinsk valley contained five freshly implanted zygotes. Her milk glands were enlarged but devoid of milk (Rukovskii, 1955). Side by side with the spring appearance of kittens, some very late litters have been found in the Caucasus. On August 3 near Gelendzhik four kittens aged 10 to 12 days were found in the hollow of a tree (Obraztsov, 1928). In the Zakatalo-Nukhinsk valley on August 15, 1950 a female contained seven fetuses with an average length of 68 mm each, indicating the last third of the gestation period. This cat would have given birth early in September. In the same region newborn kittens with a body length of 43 to 49 cm and weighing up to 2,300 g were found in early August and September (Rukovskii, 1955). The number of kittens in a litter in the Trans-Carpathians is three to six (Konyukhovich, 1953), in the Carpathians four (Lindemann, 1953; FI. Strautman), in Moldavia three to five (Lozan and Korchmar’, 1965), and in the Caucasus three to seven, most often four. Of the 12 litters studied, there were 4 kittens in seven, 5 in two, and 7 in one. In hybrid litters of forest wildcat with domestic cat, there were no more than five kittens per litter (Haltenorth, 1957). In Western Europe the litter usually consists of three or four kittens. The average for 41 litters was 3.3; one litter contained one kitten, 7 had two, 15 had three, 14 had four, and 4 had five each. In Scotland a litter contains two to four kittens (Haltenorth, 1957) and in France two to six (Beaufort, 1961). Growth, development, and molt: Kittens are born blind and helpless. They gain sight on the 9th to the 12th day. At that time they are covered with a fuzzy coat; the top of the body is very pale yellow, with dark brown spots fusing on the back into broad dark bands. The hind legs are mottled with numerous narrow stripes. Five broad dark rings occur on the tail and the tip is dark. The sides of the head, in the region of the ears and cheeks, 325 437 have a reddish tinge. A reddish spot is present on the nose. Some dark stripes traverse the top of the head and the tips of the ears are dark. The underside of the body is very pale yellow. The neck and underside of the chest are sprinkled with faint dark-colored spots. These kittens differ from domestic ones in their larger, more broadly opened ears. The weight of four newborn kittens in the Cracow Zoological Garden was 38, 42, 44, and 45 g. Growth and development of kittens of European wildcats proceed rapidly (Lindemann and Rik, 1953; from Haltenorth, 1957): Age (days) Weight (g) Age (days) Weight (g) 0 45 100 °° 1.044 i 106 130, 2) 1,654 45 308 160 2,033 52 380 190 2,841 66 543 220 2,866 80 804 250 3,128 Incisors erupt at the age of 14—30 days and milk teeth are replaced by permanent ones at the age of 160—240 days. Sexual maturity is attained in 300 days. The young begin to hunt with their mother at the age of 60 days and travel with her up to 140 to 150 days, after which they move independently. The first encounters with solitary young cats which have left their nest occur in mid-June in Caucasian preserve (Teplov, 1938). At this time they leave their mothers and hunt for food on their own. By August young cats hunting singly are a common phenomenon. Lactation continues for three to four months but kittens begin to eat meat also (mice, young hares, young rabbits, and small birds) at the age of 1.5 months. When slightly over a month old they set off from the den and play, often climbing trees. Trees soon become the favorite places for play and serve as refuges when danger is near. By the end of September in the Caucasus the young of spring litters are only slightly smaller than adult females in appearance and weigh over 2.0 kg. In Caucasian preserve a male caught on September 10, 1936, weighed 2,235 g and a female caught on November 19, 1936, weighed 2,800 g; a female on December 25, 1936, weighed 2,500 g and a female on January 22, 1937, weighed 1,900 g. As long as her kittens are small the mother is extremely restless, does not leave them alone for long, protects them from attack by small predators, and in the event of danger, transports them to a new den. The maximum longevity of the wildcat is 21 years; usually it survives for 13 to 14 years (Shortridge, 1934). The report that one cat lived for 39 years (Pidoplichko, 1956) is not reliable. Cats of the silvestris group molt twice a year. Spring molt ends in May and autumn molt in mid-November. The [winter] coat is fully developed from mid-November through March. Enemies, diseases, parasites, mortality, and competitors. Enemies of wildcats are almost unknown. In most samples of food and feces of the wolf in the Caucasus, remains of wildcats have not been found. Only one predatory battle has been witnessed. After a stubborn struggle a forest marten overcame a wildcat, tore open its abdomen, and ate its liver (Satunin, 1915). However, considering that the cat usually lives at places where there are rocks and tall trees for refuge or dense thickets and abandoned burrows, serious enemies can hardly exist. In Central Europe many young cats are killed by marten (Trauboth, 1961). The diseases suffered by wildcats have not been studied. In zoological gardens they sicken and die from infectious enteritis (Tsvetaeva, 1949). Wildcats are highly parasitized by helminths. Of ten cats examined in Caucasian preserve, eight were infected with up to 70 nematodes, and between 6 and 73 cestodes were found in all ten (Teplov, 1933). In some cats in Georgia five species of helminths have been identified: Hydatigera taeniaeformis, Diphyllobothrium mansoni, Toxocara mystax, Capillaria feliscati, and Ancylostoma caninum (Rodenaya, 1951). In Azerbaidzhan cats are infected by Hydatigera krepkogorski and T. mystax (Sadykhov, 1955). In the Trans- Caucasus the majority of cats caught were infested with the tick Ixodes ricinus. In some summers cats there are severely infested with fleas of two species: Ceratophyllus consimilis and Ceratophyllus sp. The first has been found in large numbers in Norway rats living in the forests (N.N. Rukovskii). The competitors of wildcats are jungle cat, jackal, fox, marten, and other predators. The jungle cat is a particularly serious competitor. However, the two cats usually occupy different biotopes. In the Caucasus jungle cats inhabit the lowland sections while forest wildcats reside higher up along the mountain slopes in beech forests. Usually at places where wildcats are numerous jungle cats are few or altogether absent, and vice versa. In the Lenkoran lowland jungle cats are numerous and wildcats absent. Sometimes such a distribution can be explained by failure of the ranges to meet in a small territory. Population dynamics. This aspect is almost unstudied. In the Car- pathians, following a year of a rich harvest of beechnuts, mouse-like rodents proliferate and in roughly one year’s time the cat population likewise shows a significant rise. The reverse correlation has also been noted — а decrease in cat population in years when the rodent population is low. As a result of fluctuations in the cat population the number of skins tanned annually varies more than twofold (F.I. Strautman). The cat population increases in alternate years or at intervals of two or three years. Evidently in the Caucasus, 326 326 Fig. 192. Footprints of a wildcat оп sand. Match box for relative scale. Azerbaidzhan. December, 1957. Photograph by М.М. КиКоузКи. population shifts in wildcat depend on the availability of ‘‘писе,’” which are particularly abundant after a good crop of beechnuts and acorns. The cat population shouid decrease significantly in unusually snowy and protracted winters. In such winters it becomes difficult for these predators to obtain food and they suffer from starvation (for example, in the winter of 1953/1954 in the Kuban). Cats living in the swamp forests of the Dnestr quite often die not only in spring but also in summer floods (Brauner, 1914). Floods are fatal to kittens in the swamps of the Prut, Danube, Kuban, Terek, and valleys of other rivers. Finally, the population may fluctuate as a result of epizootic diseases. In recent years in Azerbaidzhan the cat population has decreased considerably due to intensive shooting (N.N. Rukovskii). Field characteristics. Generally bigger than domestic cats. Their color is monochromatic, mottled with dark transverse bands which are sometimes barely distinct. The tail is furrier than that of domestic cats and looks chopped; there is a light-colored spot on the throat. Wildcat differs from jungle cat in variegated coloration, and a much longer, and, chiefly, in a more luxuriant tail. The tracks resemble those of domestic cats but are bigger (Fig. 192). Generally near trails leading from a wildcat’s lair broken reeds can be found, on which the cat has cleaned its claws; on the trails themselves lie relatively large-sized fecal pellets with a blunt end. This cat eats only 327 440 the internal organs of large prey: lungs, heart, and liver. Its voice is similar to that of domestic cat but coarser. (A.S.) Practical Significance A fur-bearing animal, but the skin is of little value; the average price after tanning is about 50 kopecks. Pelts play only a small role in skin-processing plants in the Trans-Caucasian and Caucasian republics. The output of tanned skins in recent years can be judged in part from the data given in Table 15. One should keep in mind, however, that tanned skins of wild and other cats are classed under the general name “‘wildcat”’ and hence only information about the catch of “‘wildcats’’ in the Ukraine and Moldavia may be attributed entirely to the forest wildcat. The All-Union standard for pelts of wildcats refers to the group ‘‘wildcat,’’ which is further subdivided into ‘‘forest’’. When living close to cultivated areas wildcat is useful in destroying rodents (see above). In most regions, however, it is a pest of game, killing nutria, muskrat, hares, pheasants, partridges, and waterfowl. In the Caucasus this cat, along with jungle cat and jackal, is considered a major hindrance in nutria husbandry. It has long been a serious pest of muskrat breeding on the Prut, Kuban, and Terek. At some places it also causes damage to poultry farming. Table 15. Tanned skins of ‘‘wildcats’’ (total) in the Soviet Union system of Zagotzhivsyr’ [raw materials procurement] Region of procurement 1948 1949 1950 1951 Ukrainian SSR 65 120 69 33 Moldavia SSR 42 15 2D 25 Azerbaidzhan SSR 2,675 4,302 3,662 3,595 Georgia SSR 505 387 394 2,183 Armenia SSR 116 Pr aS Suc 1557)" 108 Northern Ossetia 212 325 211 262 Stavropol’ 230 951 147 72 Groznensk district 1,660 1,900 1,173 792 Dagestan ASSR 443 910 817 960 Kabardino-Balkarsk ASSR 84 24 106 146 Krasnodarsk territory 3,049 2,878 1,971 1,630 Total for the Soviet Union 9,141 11,367 8,727 9,806 4SThe ‘‘запИагу”” role of this carnivore should also be taken into consideration. Ву des- troying mostly weak and diseased animals, it ‘“‘improves”’ the fitness of the game population. 328 441 Special hunts for wildcats were not practiced in the past. Many were caught in traps and other snares set for marten, and by shooting, using dogs to tree them. Following their intense persecution in Trans-Caucasus as a pest of game, they are now caught in traps placed on their pathways without a bait, or at abandoned burrows of badgers and foxes. The pelts are used in ladies’ fur coats and sometimes converted into imitation sealskin by cropping and dyeing them black, but the color of such dyed skins is more variable than in other furs. While the forest wildcat is one of the many predators in the Caucasus and Trans-Caucasus, it is now extremely rare in the western parts of the range within the Soviet Union (except some western regions of the Ukraine). In Belorussia, the Ukraine (except the Carpathians), and Moldavia, it is an interesting monument of nature and should be conserved there by banning hunting until its population is restored. Hunting wildcats is already banned in the Trans-Carpathians and other western regions of the Ukraine. The small population of this cat in Soviet forests poses very little danger to game; on the contrary the cat has become a favorite object of hunting. (A.S.) В. GROUP LIBYCA—STEPPE WILDCATS“ 1780. Felis lybica.*’ Forster. In Buffon. Naturg. vierfiiss. Thiere, vol. 6, p. 313. Gafsa, Tunisia. 1791. Felis ocreata. Gmelin. Anhang zu Bruce. J., Reisen in Africa, vol. 2, p. 27. Abyssinia. 1830. Felis ornata. Gray. Illustr. Indian Zool., vol. 1, pl. 2. India. 1834. Felis servalina. Jardine. Nat. Library, Felinae, p. 232. India. 1874. Felis caudata. Gray. Proc. Zool. Soc., London, p. 31, pl. 6. Dzhanydar’ya by southwestern part of Aral Sea (northern border of Kyzylkum). Not Kokand and not Bukhara! (Birulya, 1912). 1876. Felis shawiana.*® Blanford. Journ. As. Soc. Boreas, vol. 45, p. 49. Yarkand, Chinese Turkestan. ©Also called spotted, and long-tailed cat. ‘7Evidently a typographical error by the author. Usually the correct classical spelling (libyca) is used. 48Pocock (1939) stated that in the material used by Blanford for his description, the pelts belonged to steppe cat and the type was designated as a skull of F. сйаи5; hence the name shawiana should be placed among the synonyms of jungle cat. This view has been accepted by some zoologists (Ellerman and Morrison-Scott, 1951 and 1966). Usually (Ognev, 1935; Haltenorth, 1953; Weigel, 1961) the name shawiana is used for steppe cats of eastern Turkestan. This is the correct usage, further supported by the fact that (previously ignored) jungle cats in the east do not occur farther than Syr-Darya (see “‘Geographic Distribution’’ of this species) and have not been known for eastern Turkestan. Steppe cats are widespread there (Murzaev, 1966). The name shawiana is the first among the series of names assigned to cats of Central Asia. 442 1914. Felis caudata morpha shnitnikovi. Birula. [=] Birulya. Ann. Mus. Zool. Ac. Sc. Petrograd, vol. 19, p. 11. Semirech’e. 1914 (1915). Felis caudata griseoflawa. Zukowsky. Arch. Naturg., A 80, 9, 95. Upper course of Chu. 1914 (1915). Felis caudata longipilis. Zukowsky. Ibid., A 80, 9, 97, District south of Balkhash. Nomen praeoccupatum—F. /ongipilis Fitzinger, 1868 (tiger). 1914 (1915). Felis caudata macrothrix. Zukowsky. Ibid., A 80, 10, 127. New name for F. c. longipilis Zuk., 1914. 1914 (1915). Felis (Felis) murgabensis. Zukowsky. Ibid., A 80, 10, 127. Takhta Bazar on upper Murgab. 1914 (1915). Felis caudata matschiei. Zukowsky. Ibid., A 80, 10, 130. Foothills of Kopet-Dag at Geok-tepe west of Ashkhabad, Turkmenia.*” 1914 (1915). Felis (Felis) maimanah.° Zukowsky. Ibid., A 80, 10, 130. Maimene, northern Afghanistan. 1930. Felis ornata issikulensis.-! Ognev. Zeitschr. Е. Sdugetierk, vol. 5, р. 67. Northwest shore of Issyk-Kul [Lake]. 1945. Felis ornata araxensis. Sarkisov. Dokl. AN Arm. SSR, 3, 3, 88. ‘“Lowlands of southeastern Trans-Caucasus’’; more precisely (р. 89), steppe at Arazdayan on the Araks (about 45° E. Long.). (V.H.) Description In general appearance steppe cat is very similar to forest wildcat, but appears somewhat smaller and lighter. Tail relatively slightly longer (usually more than one-half length of body) and, more importantly, thin, and not truncated, but pointed at the tip. Comparative lighter color of steppe cat due in part ‘Reference to the type locality has been corrected. Usually it is shown as 110 versts south of Geok-tepe in the Trans-Caspian district at 38° М. Lat. and 57° 30' Е. Long. in the Kopet-Dag. The point 110 versts south of Geok-tepe falls far inside Iran, deep in the mountains, roughly midway between Geok-tepe and Sabzevar, and very far from the foothills of the Kopet-Dag lying in the immediate proximity of Geok-tepe. Latitude 38° passes right by Geok-tepe along the northern foothills of Kopet-Dag. ‘°Zukowsky bestowed this name based on a description of skins of Felis caudata from Maimene published by Scalli (1887). Ellerman and Morrison-Scott (1951 and 1966) rather uncertainly (with a question mark) assigned it to F. chaus. However, according to other researchers (Ognev, 1935; Haltenorth, 1953), it is quite clearly a description of skins of steppe cat. A skin preserved in the Calcutta Museum (India), excellently described by Dr. B. Biswas (Zoological Survey of India), has not been preserved. This zoologist also thought that the skin from Maimene belonged to a steppe cat (F. libyca). (V.H.) °!The form has been described from a feral domestic cat and from hybrids of wild F. s. caudata and domestic cats. Material in Zoological Museum, Moscow University. 443 330 to the fact that, even in winter, its coat does not attain the levels of density, 331 length, and luxuriance seen in forest wildcat. Hair on tail much shorter in the former and not so dense, luxuriant; close-fitting, hence tail appears much thinner.°* Bare section on nose not flesh-colored (pink) but black or dark (aig, 1198). ‘*Pelage not long, fairly soft, and not sparse. On the back about 3,500 hairs occur per cm, with eight of underfur to every top hair. Hair on abdomen longer than on the back (abdominal type of coat) and guard hair stands out prominently from top hair; various categories of top hair (four) do not differ greatly in length, and underfur equal in length to hair of category IV. Differences in hair thickness are considerable, however. Length of guard hair on back 51 mm and thickness 92 microns; corresponding values on abdomen 58 mm and 59 microns. Length of top hair on back 43, 39, 37, and 35 mm and thickness 102, 76, 56, and 42 microns; corresponding values on abdomen 54, 49, 46, and 44 mm, and 48, 42, 35, and 31 microns. Length and thickness of underfur on back 35 mm and 23 microns and on abdomen 38 mm and 21 microns’’ (В.Е. Tserevitinov). From the foregoing description it is obvious that the hair coat of steppe cat differs greatly from that of forest cat (Caucasian) in type of fur. The main features of coloration of the steppe cat are quite specific but its details diverse and variable. A detailed description would be quite com- plicated therefore, and only the main features are described. General background color of the skin in the upper part of the body very light (Fig. 194). In darker varieties, a fairly pure light gray color; pale yellow or sandy shade absent. In many varieties skin steadily fades and acquires a yellowish or even sometimes distinct yellow tone. When the color is highly faded, the background is a very pale white with a light creamy-yellow (pale yellow) tone. Along the middle of the back (along the spine) the main color is usually somewhat darker, forming a more dark gray, brownish, or ocherous band corresponding to the general background but varying in different forms. In some animals, mainly the extremely light colored ones, darkening on spine very faint or almost not visible. Entire upper body covered with small, round, solid spots. Their diameter in different parts of the body differs, but only insignificantly and is associated mainly with length and density of pelage in a given region. At places (back) where the coat is very short and dense the diameter of the spots is less than “It is possible that some differences in body proportions do occur between forest and steppe cats, the latter having somewhat iarger heads. 5ЗРезсирНоп based on material from Turkestan and southern Kazakhstan (about 70 winter skins in the collection of MMU). It differs significantly from the description commonly found in literature, given by 5.1. Ognev (1936). This author had only a small sample of material and exaggerated the geographic importance of some features. “AOIVUIOY “NV Аа yey ‘Авто vywpnvd siysaazis (sya) sya ‘eo э49э15 `е6т Зы 6ZE 445 330 Fig. 194. Adult male steppe cat in winter coat. Aksu-Dzhebagly preserve. Talassk Alatau, western Tien Shan. January, 1953. Photograph by F.D. Shaposhnikov. on the abdomen where the coat is longer and more sparse. This feature also affects the sharpness of the outline of a spot, but only minor individual — variations are seen. The nature of the spottedness evidently also depends on age—small-sized skins apparently appertain to newborn kittens, which have the smallest and also densest spots. - On the whole the spotted pattern is highly variable, due to various combinations of features (Fig. 195). The spots may be relatively large or small, sharply contoured or with weakly or greatly blurred outline, and disposed densely or relatively sparsely. 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> (Fig. 29). It is somewhat iarger, quite voluminous, and bulges uniformly. It mainly differs in the iarger volume of the ectotympanic chamber. As a result the section between the ear and the Eustachian openings is more roomy and the anterior boundary projects noticeably forward. It is shifted much more forward from the opening of the Eustachian tube than the boundary of the section lying inside. In the skull of forms in the silvestris group, both these boundaries fall at the same level in relation to the sagittal plane, or the boundary of the inner plane is shifted forward even slightly more. The anterior margin of the ectotympanic chamber in the steppe cat lies at the level of the anterior margin of the postglenoid process or almost at this level; in the forest wildcat it is significantly shifted posteriorly and situated behind the posterior margin of the postglenoid process. The opening of the external auditory meatus in the steppe cat is large in absolute and relative terms, its size increasing in relation to the increase of the ectotympanic chamber. The relatively large size of the external auditory meatus is not obvious because of the overall size of the auditory bulla (profile); furthermore, in the steppe cat the maximum diameter of this opening is greater than the length of the crown of the third upper premolar; in the forest wildcat it is smaller or equal to it. Such a structure of the auditory bulla of the steppe cat represents a distinct adaptation to desert conditions, totally homologous to such adaptation in many other mammals (carnivores and rodents). There are no differences in dentition but the peak on the anterior- SAs pointed out above, some workers (Pocock, 1951) assume that there are no dif- ferences between the skulls of F. silvestris and ‘‘F. ПБуса,`` and that these *‘species’’ differ only in color. Other characteristics have been verified on a large number of animals in both groups, but only from within the Soviet Union. Evidently these characteristics are common to animals of adjoining territories. Material from Africa has not, however, been investigated. The form sarda (‘‘transitional’’) has auditory bulla of the forest wildcat type. 334 451 Fig. 197. Young male steppe cat. Badkhyz preserve. Southern Turkmenia. May, 1962. Photograph by Yu.K. Gorelov. inner cusp of the upper carnassial tooth is evidently somewhat better devel- oped and sharper. Both upper premolars were present in 66% of the animals examined (in 30 of 45 males and females taken together), both were absent in 18% (8 ef 45), or one absent in 15 to 16% (7 of 45). In males (22) both teeth were present in roughly 64% (14), both absent т 27% (6), and one absent in 9.0% (2). In females (12) both premoiars were present in 66% (8), both absent in 8.0% (1), and one absent in 25% (3) (material from Turkestan). These ratios are somewhat different than the corresponding ones in the Caucasian forest wildcat. Sex-related and age-related skull changes correspond to similar changes in the forest wildcat. Geographic variation has been noticed but not thoroughly studied. It is not seen within the boundaries of the Soviet Union. Tn body and skull dimensions forms representing the group correspond, on the whole, to forest wildcats of Europe and the Caucasus but. on the average are smaller; some southern animals are significantly smaller. in any case the maximum size and weight atiained by forest wildcats of Europe 336 452 are not attained even by members of the biggest subspecies of steppe cats.°° This pertains also to the Turkestan form (see ‘Geographic Variation’’). Females are perceptibly smaller and lighter than males. Within the species as a whole Bergmann’s rule is distinctly manifested (biggest cats found in Europe and Middle Asia); the same is true of Gloger’s rule—most richly colored cats found in Europe and the Caucasus among them especially in Scotland—the whole of the desert-steppe members of the [Буса group are pale; some desert forms of Africa are particularly pale and exhibit a much reduced spotted pattern. Allen’s rule is manifested in body proportions. These rules often apply within each group of subspecies individually. (V.H.) Origin of Domestic Cats At present it is thought (Pocock, 1951; Haltenorth, 1953 and 1957; Zeuner, 1957) that the ancestral form of domestic cat was the North African bay or Libyan cat, F. (F.) s. libyca Forst. It was domesticated in Egypt very long ago. According to some authors the first reference to domestic cat dates back to about 4000 B.C., or to the First Dynasty, about 3000 B.C. By 2000 В.С. at the beginning of the Middle Kingdom, cats were already quite common in Egypt as domestic animals and widely distributed. According to other authors (Zeuner, 1967) portrayals of cats in the Middle Kingdom represent wild animals and it was only in the New Kingdom, about 1600 В.С., that the cat becante a domestic animal, i.e., domestication occurred not very long ago.” Initially domestication of cats arose ‘from the need to combat house and granary rodents. Mystic or religious importance was assigned to the cat later. It was even considered a sacred animal (cult of goddess Bast at Bubastes). Bodies of cats were mummified and preserved in large numbers. A study of a large number of these mummies (Morrison-Scott, 1952) revealed °°This refers to normal variation. As pointed out above, especially large wildcats of western Europe (weighing from 11—15 to 18 kg), if these figures are not erroneous ог exaggerations, probably represent hybrids with domestic cat, i.e., an exotypical phenomenon like the *tgiant’’ Amur cats. Hybrids of steppe and domestic cats do not attain such “‘giant”” sizes; at least they are not known to occur within the Soviet Union. ‘7B. Brentjes (1965) assumes that the cat was domesticated in the Near East much earlier than in Egypt. According to him, representations of cats in Erihon and the Jordan valley, from the end of 6000 and early 5000 B.C., depict domestic animals, as do representations from southern Iraq from early 300 B.C. Statuettes of cats from Kachilar in Asia Minor date to 6000 B.C. Thus the Libyan cat, F. 5. libyca, is not the only ancestor of domestic cat; it had already hybridized with the Middle East cat. This view has found no confirmation (Zeuner, 1967). Brentjes has not taken into consideration that forest wildcats lived in the Kachilar region, which removes the entire question of ancestry to a different plane. (V.H.) 453 that F. 5. libyca was domesticated and, furthermore, long before our time the Egyptians had a special domestic form of cat, which was slightly bigger than the wild one (‘‘F. bubastis’’ Hempr. and Ehrenb., 1832). The possibility should not be completely excluded of other pockets of domestication in antiquity, independent of Egypt, in the agrarian states of the Near and Middle East, and India, 1.е., in the range of the subspecies of group ornata—caudata, primarily in Persia and Mesopotamia. However, such a probability appears dubious and evidence quite inadequate. In these countries the cat more likely arrived from not-so-distant countries such as Egypt. Ancient China could not have produced the domestic cat because the species silvestris—libyca was absent there. The domestic cat of China came from the Middle East through India just before our era (not earlier than the sixth century). Europe was also not the home of cat domestication and the form F. s. silvestris cannot be included among the ancestors of domestic cats. Not only the low population density and lower cultural level of the inhabitants of Europe, but also the wild nature of the forest cat, exclude this possibility. The forest cat does not tame as readily as the steppe cat and is far less suited to domestication. The cat in Europe either came from Egypt directly or via the Near East. Evidently there were two routes of range expansion—via Italy (Rome) or via the Balkans and Black Sea region. The cat appeared first in Europe in the Black Sea region (Ol ’viya, sixth to the first century B.C.; Pidoplichko, 1952). Reports of its appearance in Rome are dated much later; the first reference pertains to the fourth century A.D. The cat colonized central and western Europe only in the Middle Ages and was a rarity in the far west (England) even in the tenth century. Colonization of cats in Russian territory occurred roughly in the same epoch as in central Europe. As indicated by the period in which some finds are reported and the overall political and cultural situation obtaining in ancient Russia (relations with Byzantium in particular), the cat evidently came into Russia not only from the west but also from the south. Its remains are known from the seventh and eighth centuries A.D. in the Sum district (Volyntsovo Putivl’sk region), from the eleventh to the thirteenth centuries in the Zhitomir district, and from the twelfth and thirteenth centuries in Kiev and elsewhere. To the north remains of domestic cats have been found in horizons of the seventh to the ninth centuries in Staraya Ladoga, tenth to the thirteenth centuries in Moscow, eleventh century in Novgorod, and thirteenth and fourteenth centuries in Pskov and elsewhere (Pidoplichko, 1956; Tsalkin, 1956). As mentioned earlier, while discussing hybridization of domestic and forest wildcats, a change in landscape and a decrease in population of forest cats could have led to instances of hybridization with domestic cat. However, 337 454 the influence of domestic cat on wild was greatly exaggerated until just recently. It was thought, and sometimes still is, that domestic cats exerted a major influence on wildcats. The picture in fact was otherwise. Wild forest cats evidently exerted some influence on European domestic cats. Animals with a tiger-like coat (transverse stripes or distinct vertical rows of spcts on the body) exhibit a pattern characteristic of forest cats of Europe. Even in Turkestan this pattern is found to varying degrees only in hybrids of steppe and domestic cat and not seen among ““pure’’ wildcats (Е. 5. caudata). in general, however, the influence of European forest cat on domestic cat was very insignificant. It has been suggested that the Angora cat (also called Persian cat in the west) evolved as a result of hybridization of domestic cat with the Caucasian forest cat in Asia Minor. Its long coat was apparently acquired from wildcat together with some other features of its ““face’’ and general appearance (Pocock, 1907; Schwangart, 1932; and other earlier authors; Haltenorth, 1953). This suggestion has not been confirmed. The lone argument is based on the assumption that this form originated in Asia Minor (Angora-Ankara) where the forest wildcat is present. Such a view is hardly justified; long-haired breeds of domestic cats are also known in Middle Asia (‘‘Bukhara’’), Siberia (‘“Siberian’’), and China. Evidently these species are also products of selection like other long-haired “‘Angora’’ animals (rabbit, goat, and guinea pig) and Pekinese dogs. In discussions on the origin of domestic cats certain other views have been expressed which are no longer accepted. Thus, even Pallas (1811) suggested (some authors have interpreted his suggestion as an affirmation) that the Angora cat is derived from the manul cat since “‘domesticated manul cats readily mate’’ with domestic cats (“‘licet mansueta cum hoc facile coeat’’). Pailas’ suggestion had to await confirmation for want of anatomical data. Such data, now available, clearly reveal that the two species are so far apart from each other that there is no justification whatsoever to assume any contribution of the manul cat to the evolution of domestic cats. Strangely, this question is still raised by some authors even today. As an extreme rarity, under artificial conditions, production of some hybrids 1$ possible, but such a rare hybrid could not have influenced the evolution of domestic forms. Furthermore, hybrids of this type are not known, although instances of mating probably do occur (see section below on Pallas’ cat). In India it was believed that the domestic cat often mated in nature with the jungle cat. Among British cat lovers and breeders there is (was ?) even a belief that one of the color types of domestic cats (‘‘Abyssinian’’) was produced by an admixture of the blood of jungle cat. This view 15 erroneous (Pocock, 1939). Some rare instances of hybridization of jungle cat with domestic cat in nature cannot be ruled out. However, reports on this 338 455 phenomenon as well as on the resultant hybrids are evidently not available for India. There are also no Soviet data on this subject. Hybrid kittens have been bor in captivity (London; Gray, 1954) to domestic and jungle cats. Information that the Egyptians extensively encouraged the mating of jungle cat with a domestic counterpart and used the resultant hybrids for hunting (E. Bogdanov, 1913) is erroneous, and does not figure in recent literature (Brentjes, 1965; Zeuner, 1967).8 In any case, among the 190 skulls of mummies of cats from Gizekh (Giza, 600 to 200 B.C.) which were thoroughly studied (Morrison-Scott, 1952), only 3 were found to be skulls of jungle cat.°° The rest (187) were domestic cats (‘‘F. bubastis’’) or wild local bay cat, F. 5. libyca. Probably the Egyptians used for hunting (if the practice actually existed) tamed and trained jungle or domestic cats. In any case, there is no question of the contribution of jungle cat to the evolution of domestic cat; this possibility has been eliminated. Thus the only true ‘‘primary’’ ancestor of domestic cat, “РЕ. catus,”’ is the species Е. silvestris and the Egyptian subspecies of the group of steppe cats, F. 5. libyca. The European forest cat, F. 5. silvestris, could only be of secondary importance in the evolution of the European domestic species and some of their characteristics. The rest of the subspecies probably influenced to some extent its characteristics in different parts of the range in Asia and Africa, as a result of limited hybridization with the fully developed domestic form. However, this influence is not relevant to the evolution of the domestic cat. The modern interpretation of the species F. si/vestris and other available information have proved the origin of the domestic cat from two species (V.H.). Systematic Position The great morphological similarity, the existence of subspecies with inter- mediate features, strict vicariance of both groups and individual subspecies, and free hybridization of forest and domestic cats in nature, support the notion that forest and steppe cats are a single species (or supraspecies). F’. silvestris as a species, on the whole, is very closely related to the jungle cat. In the “Egyptian depictions of cats ‘on the hunt”’ do not permit a differentiation of domestic or bay cat from jungle cat; trying to distinguish hybrids of jungle cat with domestic cat is well- nigh impossible (V.G. Heptner; В. Вгепцез). °Haltenorth (1953) states that there were 12. Some misunderstanding occurs here; of the 12 listed by Haltenorth, 9 are recent skulls from the collection of the British Museum, which served as comparative material for identification. Thus the percentage of jungle cats in the total number is not 6.36 but only 1.58. 456 composition of the Soviet fauna this represents а progressive stage in the course of specialization of the feline type from Amur cat to manul cat. However, the morphological and biological isolation of the two species is very sharp.© In the series of Soviet species of the genus, Е. silvestris s. [. occupies a somewhat central position between the less specialized species F. euptilura and F. chaus, just as does the side branch formed by F. caracal and Р. lynx, and the highly specialized Е. margarita and Е. manul, which stand at the end of the “‘direct line’’ of specialization. At the same time the species is not entirely monolithic and within those limits the silvestris s. str. group is very close to the jungle cat. Subspecies of the ПБуса group exhibit features of further development (specialization) of the feline type. This is evident from the structure of the auditory bulla. The ectotympanum is somewhat enlarged, and this is an insignificant but fairly definite step forward toward the development of the type of bulla seen in more specialized cats such as the sand cat and manul cat. It is signi- ficant that this line of separation divides a single species into two groups of subspecies. (V.H.) Geographic Distribution These cats are found in the open steppes and deserts, and in part montane regions of Africa, the Near East, parts of Central Asia, and northern India. Geographic Range in the Soviet Union The range in the Soviet Union includes the extreme northern part of the range of the species and covers a relatively small southern part of the country. It consists of two individual sections—a small one in the Trans-Caucasus and an extensive section in Middle Asia and Kazakhstan. These two sections join in the south outside the borders of the Soviet Union. In the Caucasus steppe cats occur only in the extreme south and south- east: in the Lenkoran lowland (evidently also on the Talysh); steppes of the lower Kur’, i.e., in the Mugan and Salyan steppes; in the Araks valley and the adjoining slopes within Nakhichevan Autonomous Soviet Socialist Republic and Armenia; i.e., west of 46° 30' Е. Long. In Armenia steppe cats are known from Arazdayan at the boundary with Nakhichevan district, Daval on the Araks, and the slopes of the Saraibulag range in Vedinsk region; 1.е., Definitive recent data are not available about natural hybrids between them, even through domestic cat, although extensive geographic and biotic contacts do occur. At the same time instances are known of natural hybrids with Amur cat through domestic cat, the former being a distant species (see ‘‘Origin of Domestic Cats”’ and *‘Systematic Position’’ of Amur cat). 339 340 457 in the district east and southeast of Yerevan. They possibly reach Leninakan along the Araks. In Middle Asia steppe cats are widely distributed. In the extreme north- west the range encompasses the deserts of the Pri-Caspian plains between the lower reaches of the Volga and Ural, in the west almost to Volga (Dzhambai), in the north including the Dzhangalinsk region of Western Kazakhstan district (region of Kamysh-Samarsk lakes), and in the south to the northern coast of the Caspian Sea. Farther east the northern boundary of the range runs through the middle course of the Emba, encompassing southern Mugodzhar, and crosses roughly through the city of Irgiz, passing by Lake Chelkar-Teniz to the north. From there it runs in to the Ulutau mountains, later descends somewhat to the upper reaches of the Sarysu, and encompasses to the north the Kazakh foothills, but does not reach Karaganda. Even more eastward it ascends again slightly northward to Karkaralinsk. Later, the northern boundary descends south, passing slightly south of Semipalatinsk, and encompasses the Chingiztau and the Zaisan depression with the Chernyi [Black] Irtysh.. There, the boundary runs into China. _ The boundary as outlined in Kazakhstan representing the sum total of all available data, in some parts evidently making for an exaggerated picture of the actual distribution of the species in the north (Fig. 198). The point is that at some places the boundary is probably based on rare intrusions, at others evidently on imported skins, and in several sections, particularly the entire region north of Balkhash, data possibly confused with that for manul cat (especially local information). The line describing the region of permanent habitation of the steppe cat, as drawn from recently collected data of the mid-1960’s, evidently runs much more to the south. It proceeds from the mouth of the Emba to the northern end of the Aral Sea, then to the mouth of the Sarysu, encompassing the lower half of the river, runs on to the Chu River, and encompassing the southern part of the Bet-Pak-Dala [desert], reaches the western shore of [lake] Balkhash. Subsequently it proceeds along the southern shore of the lake, and arcing around the Alakul’ depression, passes into China. The occurrence of the steppe cat in the Tarbagatai region and Zaisan depression, as also in the Volga-Ural plain, remains dubious (A.A. Sludskii). For all of that, there is evidently no foundation to call into question or to contradict the significance of data for earlier dates in our century. There is no mention of factors leading to a shift in the range during recent decades. It is highly possible that in a series of sections the boundary occupies an intermediate position between the two lines. To the south steppe cats are widely distributed. In the region west of the Amu-Darya they cross the state boundary everywhere and are encoun- ‘зэщ9энН “OA ‘те\у-ишеа рие ueys vary oy) Jo зрие|ч8 oy) ит |еиние $143 JO эоцэзае oY) эзеотри! 1110$ э4} и! э5041 рие ‘(иузри|$ У‘) Neely узле8ииц2 sy} Jo цо!8э1 эц и! попеиаеч зподе вер элниуэр JO ээцэзае эзопэр 1565 ay} Ul gsou} чеёзат, рие 7181 JO sayoral Jamo] 24} UI иоцейаец Азелодитэ} эзопэр YIOU JY) UI зулеш поцзэпо ‘(иузри]с “y'V) роцэ4 swoAy[nw в 1эло SUIYS рэцие; Зширле8эл ипоцеизозит рэшациоо UO pose э8из1 Jo Алерипо4 изэцуиои —< 480 э943} Jo surys рэплодии Jo (997 “31,4 295) Ц эаеаолА Ajysry si 3] ‘элизелэ ит э1аетеле вер до [2101 Wns UO рэзеа Алерипоа—Т 22 [NUL иээлм1эа YSINSUYSIP JOU ртр вер эшоз Jv ‘(WY ит э[е2$) UOIUA 191406 э4} UI вррирд siysaazis (51]2>]) Say ‘eo э94э1$ Jo э8иел Jo Алерипо4 изэчуом ‘8618 i il | ll | Ss — 5 = Y 65 459 tered also in the Great Balkhan and Kopet-Dag, including their highest sections (up to 2,000 m above sea level). The southern boundary of distribution east of the Amu-Darya is not quite clear. Thus this species is known in the Tarbagatai, Dzhungarsk Alatau, Trans- Ш Alatau, Kirgiz Alatau (Aleksandrov range, northern slope), the Issyk- Kul’ depression and Chu valley, the Talassk Alatau and western part of the Tien Shan along the upper Arysa, Chirchik, and Keles with its tributaries, and in the Ferghana valley and its surrounding slopes. There is no doubting its occurrence along the foothills of the Turkestan and Zeravshan ranges, although there is no direct proof; it is also known from the Zeravshan valley. It lives in the Baisunsk mountains and the Kugitangtau, along the Surkhan- Darya and the surrounding mountains (Babatag), in the Gissar range along the Pyandzha and the rivers flowing into it, and in the low mountains and foothilis dividing them. Eastward the range extends there roughly to the Kulyab meridian and evidently even farther.°! Steppe cats occur throughout the plains and occupy the foothills as well as the lower and middle zones of all mountains in Middle Asia. At places, along valleys and gorges, they are found deep in the mountains and ascend to the upper limit of trees. In Semirech’e they are known from heights of 1,800 to 2,000 [m] and in southern Tadzhikistan up to 2,500 m above sea level. It is evident that in significant districts of the high montane region, such as the depths of the Tien Shan, or corresponding regions in the Pamir- Altai system, this species is absent. However, precise data about this are scanty and, in principle, all the actual positive information pertains to the territories shown above. At the same time, there is reason to believe that the steppe cat may occur, though rarely, perhaps only as an intruder, even at great heights deep inside extensive montane massifs. Reportedly it occurs *‘throughout Kirgizia’’ (О. Dement’ev, 1938), and also another cat (‘‘Felis sp.’’), possibly a different species (manul cat ?), is found in the southern Pamir at Yashil’kul’ (3,750 т above sea level) (Meklenburtsev, 1936). Thus the precise southern range in the eastern half of Middle Asia is not yet clear. Geographic Range outside the Soviet Union The range outside the Soviet Union (see Fig. 181) is unusually extensive and, if we exclude Central Asia, is very similar to the range of caracal and cheetah. Steppe cats live in Dzhungaria and Kashgaria, on the east to Kham, Lobnor, and the Kumtag sands, and in the southern and southwestern parts of the Mongolian People’s Republic (Trans-Altai and southern Gobi) and $1 Вапое based on data of Ognev, 1935; Shnitnikov, 1936; Vereshchagin, 1940, 1947, and 1958; Sarkisov, 1945; Kuznetsov, 1948 and 1948a; Sludskii, 1953; Dal’, 1954; Afanas’ev, 1960; Ishunin, 1961; and others; and original material of A.A. Sludskii and V.G. Heptner. 341 460 Inner Mongolia (Aiashan and Ordos to Yulin-fu). They also live т Afghanis- tan, Iran, Baluchistan, and India. In the latter country their range covers Punjab, Sind, Kutch, Rajasthan, and central India, probably not south of 20°. They likewise inhabit Iraq, Syria, Palestine, and the Arabian Peninsula. In Africa they are encountered throughout the mainland except the central parts of the Sahara and the western African forest region. Some authors assign the cats of the Mediterranean islands and Italy to this group (ПБуса, see above). It has been reported that the steppe cat inhabited during the Pleistocene and Holocene limits beyond its present-day range, 1.е., Ireland, Belgium, France, Spain, and Italy. In Ireland it was found along with the remains of domestic animals (Pidoplichko, 1951; and others). However, as mentioned above, the ‘‘feline’’ remains on which these conclusions are based, in all probability are those of the forest wildcat (silvestris group). (V.H.) Geographic Variation Steppe cats represent a group of forms which have been assigned some 50 names—the largest number of names for any cat. Leaving aside the un- scientific approach distinctly manifested among some authors while describing ‘‘new’’ forms, another reason for this proliferation of names may be found in the extreme variability of coloration throughout their range as a whole, as well as in each individual section. This, together with the inadequacy of collections, has created favorable grounds for an extremely narrow interpretation of the subspecies until recently. Even at the end of the 1940’s (Pocock, 1951) the species silvestris was thought to consist of two species —si/vestris and libyca—with about 30 subspecies. Ten names were assigned to forms present within the Soviet Union and four forms recognized (Ognev, 1935; Pocock, 1951). In reality the number of forms is far less than the 14 which have been suggested (Haltenorth, 1953 and 1957; for the species as a whole, 20)53. Only one subspecies occurs within the Soviet Union. Turkestan steppe cat, F. (F.) 5. caudata Gray, 1874 (syn. shnitnikovi, griseoflava, longipilis, macrothrix, murgabensis, matschiel, maimanah, issikulensis, and ? araxensis). °*Zukowsky (see synonyms) has described five *‘species’’ from a single region (flat desert zone of Turkestan and Kazakhstan) not isolated by any barrier from one another. The type localities (not the boundaries of their range!) of these **species’’ are situated at distances of no more than 150 to 700 km from each other, or from forms described earlier, or directly concordant with them. °3Weigel (1961) has enumerated 20 forms in the group under description (27 in the species as a whole), but this enumeration does not constitute a taxonomic revision. 342 461 Main coat color light, ocherous-gray; dark spots well developed through- out trunk, small, and quite sharp; narrow, dark-colored band absent along back but chain of spots present. Hair at end of tail somewhat shortened and tail pointed. Ectotympanic chamber of auditory bulla in front of auditory meatus slightly inflated and shifted forward. Maximum diameter of auditory meatus more than length of crown of third upper premolar. The description given above pertains to this form. Body measurements and weights are given in Tables 16 and 17. Body measurements of 12 adult males and 5 females from the Karshinsk steppe (south of Samarkand) are as follows: body length 530-570 and 490-560 mm, tail length 310-340 and 230-300 mm, length of hind foot 128-142 and 124-136 mm, and height of ear 60-65 and 58—65 mm (Salikhbaev et al., 1967). Weight of heart and heart index for three males from the upper Amu- Darya as follows: 25 g and 7.74%o, 30 g and 5.66%o, and 29 g and 7.25%o; corresponding values for two females: 32 с and 1.52%o and 28 g and 11.91%o. Length of intestine and its ratio to body length of the same three males and two females: 1,160 cm and 1 : 2.8; 1,530 cm and 1 : 2.7; 1,300 cm and 1:2.4; 1,190 cm and 1:2.4; and 1,400 cm and 1:2.7 (calculated from material of Chernyshev, 1958). Found in southern Kazakhstan and ? Trans-Caucasus. Outside the Soviet Union F. (F.) s. caudata occurs in adjacent parts of Iran and Afghanistan; and ? Dzhungaria. In body measurements and weight the steppe cat is perceptibly smaller than the Caucasian, although it is entirely possible that such a situation is due to random selection of material of the Caucasian form. This assumption is supported by the fact that the skull dimensions of both forms are identical. According to existing figures for the steppe cat, the head is relatively large (maximum length of skull about 18% of body length and in the Caucasian form about 15%). The tail length of the steppe cat is relatively greater — on the average 54% of body length versus 49% in the Caucasian form. Throughout the entire extent of the range in the Soviet Union dimensions of steppe cats are identical (series of skulls from Turkmenia and Semirech’e are totally identical). Similarly, there are no essential differences in color in animals from different parts of the range. A study of a series of skins (over 70) revealed that reddish types (general hue and pattern characteristics) are geographically not localized and the separation of various geographic forms, except for caudata, not justifiable. Specimens of issikulensis, described from Issyk-Kul’ (Tien Shan), which differ sharply in their dark and gray coat color and large, coarse patches (these patches have a tendency to form vertical rows), are evidently hybrids of steppe cat and domestic cat with a tiger-like coat (the type is hybrid; "9 YOoIWaS рие вАле-пшу 1э44п oyj Jo ulseq рив ешэшута, ол) Арте $иэщ1934© '590110$ 12410 рие ‘(866Т) лэЧзАшэцо `1' A “(ззоцэюс Jo AWapeoy ‘штэзпи 1е180007 “SEGT) ^эи80 `[$ JO Зиэшолизеош рие ‘иззри[с “WY Jo пензе ‘Аизлэлииг 40950 “WUNssNA] [2180[007, ay) Jo цОоПЭоЦоЗ UO poseg :эюм 9/9°< 0°ТФ SrO'Z IT reo’ 000°9 S66 T 8Т 481Э М 95 OL OV va! TL 06 gs ST sive JO 1481эН Let OST Отт oT EEE OST OOT SZ Sue] 100} pury C87 Ove а 81 ПЕ 09 OST 9% yysuay [Ie], 055 от9 Orr ST SLS 07 06 LZ Чиэ1 Apog W хеш wu и И хеш ее и == $эхэри] $21297 дару зэтеш ииру це} эута т, Jo ‘27ортво °5 (°J) *Я ‘syed э4@з1$ jo (8) з48эм рие (ии) зиэшолибваш АроЯ ‘OT э1ает, Ive 462 463 ‘OT э1ает, лэрип рэ1$| jy) sv эшез [ее] :310м ELE 0'Or УС GOL СЕТ =0т 65 4100} Гетбзеилео 1э44п до ц38иэ'] Le €LZ С TEE 196 £62 OF MOI 4300} Jaddn jo Ч13иэ] TOE 0`0 УС Lee 8'86 OTE 8Е UIPIM [еиато1$04 105 Vol УС [6Т €€Z rol Se YIPIM пеазолоуи] €OL 5'65 TZ Pu, €78 L'€9 9€ YIPIM эцешо8А7 0°16 $7'91 < [776 Z'EOL 158 LE Ч 8иэт [eseqo|Apuod € 001 €¢8 [га У’ соТ Ес 856 9Е 43843] 15912219) хеш ии u W хеш ull u ae SOXOPU] 59[241э] [NPY 5эеш пиру ие] SAYIN], JO ‘vywpnys °5 (J) yy ‘syed addajs Jo (WU) ззионтолазвош [[NYAS °/Т ЭТ, (443 343 464 cotype—evidently a feral domestic cat; Zoological Museum, Moscow University). Animals similar to those from Issyk-Kul’ have been seen in the environs of Alma-Ata (Vernovo; Zoological Museum, Moscow University), and along the Ш River (Dzhil’turanga), 1.е., from the region of occurrence of true caudata.“ Specimens from Issyk-Kul’ and other hybrids from Semirech’e have the skull features of F. s. caudata. Material from the Trans-Caucasus has not been studied and the systematic position of these animals, in spite of a description of form araxensis, is not known. The relations of the form caudata to neighboring subspecies of the libyca group are not understood. Forms from Inner Asia, at least kozlovi and schawiana*, are very close to it and sometimes even placed among its synonyms; this point of view, as far as the latter form is concerned (i.e., schawiana*), is hardly justifiable. The Indian ornata (Pocock, 1939) and the Mesopotamian vesterovi are also related to it. Compared with ornata, the Soviet form is evidently larger. * * ok The picture of geographic variation of the group is complicated by the fact that everywhere, or almost everywhere, it forms a cline. In recent decades a more realistic approach to the question has emerged, and the tendency now is toward a significant reduction in number of forms. Nevertheless, a system of strong splitting is still quite prevalent. Outside the Soviet Union, within the confines of the group, leaving aside the above-mentioned ‘‘transitional’’ forms from the Mediterranean (see р. 423), and forms sometimes assigned here but reckoned by most zoologists to be F. (F.) bieti M.-E., 1892 (chutuchta and vellerosa), the following subspecies (or groups of subspecies ?) may be considered (according to Haltenorth, 1953 and 1957, with modifications): 9) F. (Е.) 5. chawiana Blanf., 1879** (including kozlovi Satunin, 1905)—eastern Turkestan and ? Dzhungaria; 10) F. (F.) 5. ornata Gray, 1823—central and northwest India and Pakistan; 11) F. (F.) $. nesterovi Вища, 1916 (including тая Cheesm., 1920)—Mesopotamia, southwestern Iran, northwestern part of Arabian Peninsula, Syria, and Palestine; 12) F. (F.) 5. libyca Forster, 1780—North Africa from Morocco to Egypt, Nubia, former Anglo-Egyptian Sudan in the east to the Red Sea (Suakin and Massava), western part of Sinai Peninsula (Ellerman and Morrison-Scott, 1966); 13) F. (F.) s. ocreata Gmelin, 1791 —Ethiopia; 14) F. (F.) $. brockmani Pocock, 1944—Somali; 15) F. “Specimens of F. 5. matschiei, from below Astrabad, illustrated by T. Haltenorth (1953; Figs. 58 and 59), greatly resemble a hybrid of this type. A feral domestic cat is known from below Dzharkent in Semirech’e (Pocock, 1951). *Sic; should be shawiana—‘Sci. Ed. ** Sic; should be shawiana Blanf., 1876—Sci. Ed. 344 465 (Е.) 5. rubida Schwann, 1904—East Africa, southern Sudan, northeastern part of Congo basin; 16) Е. (F.) 5. cafra Desm., 1820—southern and southeastern Africa; 17) F. (F.) s. griselda Thos., 1926—central and southern Angola, northern South East Africa, and Kalahari; 18) F. (F.) mellandi Schwann, 1904—northern Angola, southern part of Congo basin, and northern Rhodesia; and 19) F. (F.) lowei Pocock, 1944—-southern Senegal, eastern Sudan, and northern Cameroon. The number of forms has been reduced in this list but in the case of Africa at least the number is still somewhat exaggerated. All of the previously listed forms are usually classified into two groups—steppe cats of the ornata—caudata group and bay cats of the ornata—libyca group. Subspecies from China, the Soviet Union, Afghanistan, Pakistan, India, Iran, and Iraq (Nos. 9 to 11 of the above list) fall into the first group. The coloration of the second group is generally paler with a well-developed pattern of spots and bands or, on the contrary, a faded pattern, almost monochromatic, and generally paler than steppe cats. All the remaining forms (Nos. 12 to 19) fall into this group. A transition between the two groups is found in the Near East and the Arabian Peninsula. Within the species in general a third group exists, represented by forest wildcats of Europe and the Caucasus, as well as a fourth group for Mediterranean ‘‘transitional’’ forms (island group; Weigel, 1961; р. 423). It is quite clear that, based on the sum total of characteristics, the group forest wildcat should not be equated with either of the above-mentioned groups (steppe and bay); this group corresponds to these two taken together. On the whole, however, such groups are of no importance from nomenclatural point of view and constitute a mere terminological convenience. (V.H.) Biology Population. In the Trans-Caucasus the steppe cat is extremely rare and caught singly. In southwestern Turkmenia they are quite common, and at places in the foothills and along river valleys are abundant; they are rarer, however, in the Karakum desert. In some regions in the west of the republics, for example in Kyzyl-Atreksk, some 200 to 400 cats are caught in a season. They are parti- cularly abundant in the Tedzhen and Murgab basins in southern Turkmenia, but rarer to the north. From 1954 through 1958 in Turkmenia 2,369 to 3,430 skins were tanned annually. In Tadzhikistan the spotted cat is quite common on the plains, near rivers, and in the foothills and particularly numerous in the Kashkakum desert. Up to 500 skins are tanned annually in this republic. In Uzbekistan the steppe cat is common in the valleys of all rivers running through the plains, and numerous in the Amu-Darya delta. In the southern part of Kyzylkum adjoining the Amu-Darya, and in the river valleys, 466 Fig. 199. Dzhelturanga in lower reaches of Ili River. Biotope of steppe cat. Tolai hare (Lepus tolai), roe deer, pheasant, and formerly tiger also lived here (as well as in sites depicted in Figs. 200 and 201). Semirech’e. July, 1960. Photograph by A.A. Sludskii. up to 2,000 cats are caught annually. They are common in the northern parts of the Karakum. In Kirgizia the spotted (steppe) cat is common in all the low-lying valleys and numerous in Issyk-Kul’ depression, 'as well as in the Chu, Kochkar, and Talas valleys. In the 1930’s up to 1,000 cats were caught in Kirgizia in a season; only 150 to 300 were caught in the 1960’s. In western Kazakhstan the steppe cat is very rare between the Emba and Ural Rivers. Their occurrence has not been confirmed in the Ural— Volga interfluve in recent decades. In places they are quite common in Mangyshlak and Ustyurt; until quite recently some 200 cats were caught in these regions in a season. They are numerous in desert regions adjoin- ing Syr-Darya, Chu, Talas, Ili, Karatal, Aksu, and Lepsa valleys. They are frequently sighted in the lower reaches of the Sarysu but rarely in the sands of its left bank (Muyunkumy, Dzhetykonur, and Dzhidelikonur). In river valleys running through the desert a hunter will normally catch 2 to 15 cats in a season. In some regions this predator was harvested in the hundreds; formerly up to 1,000 skins were tanned in a season in the Ш delta. In the Dar’ yalyk and Bet-Pak-Dala deserts steppe cats were abundant only in the extreme south, and absent in the central and northern parts of 345 467 these deserts. Today this cat is also absent in northern Pri-Balkhash and very rare in the Alakul’ and Zaisan depressions, where only single individuals are caught. In the southern half of Kazakhstan in the 1930’s an average of 8,000 cats were caught in a season; in 1931 alone.14,493 animals were caught. Now, however (1960’s), 3,000 to 4,000 cats are caught in a season. The number of skins tanned has declined everywhere, not because of a reduction in the cat population, but because of poor organization of the fur industry. In countries adjoining the Soviet Union steppe cats live almost everywhere (for example, in Iran) but are more or less common only in certain areas (Misonne, 1959); they are common in regions adjoining the Atrek and Tedzhen Rivers. In the northern parts of Afghanistan bordering the Soviet Union—Badkhyz, Karabil’, and along the Amu-Darya and Pyandzh rivers —they are even more numerous. Southward, in the montane regions, this cat is absent. In the east, in the northwestern parts of China, for example Sinkiang, steppe cats are common, but in the Mongolian People’s Republic quite rare. They are more common there only in the southern Fig. 200. Turanga trees (Populus diversifolia) and tamarisk thickets in solonchak sections with reeds in lower reaches of Ш river (Dzhel’turanga). Typical habitat of steppe cat. Semirech’e. July, 1960. Photograph by A.A. Sludskii. 346 468 regions, in the sands of the Trans-Altai Gobi, in Noyan-Somon, Khatsar- Usuni, and others. Thus in the Mongolian People’s Republic only some 20 to 30 steppe cats are caught annually (Bannikov, 1954). Habitat.© Steppe cats inhabit different types of deserts and are usually encountered in summer close to water sources, not more than a few kilo- meters away from one. They also live in desert sections of foothills, and as a rule, do not ascend high into mountains. In the Trans-Caucasus these cats live in tugais, deserts, semidesert sections here called “‘steppes,’’ and in foothills along gorges where shrubs grow at heights of 700 to 1,750 m above sea level (Dal’, 1954). In Turkmenia, Tadzhikistan, and Uzbekistan these cats confine them- selves to tamarisk (dzhingil), saxaul, saltwort, and other shrubs, and also to sparse reeds in argillaceous sections of deserts near rivers, irrigation ditches, and lakes. They are rarer in sand massifs with a good growth of herbaceous and shrub vegetation. In southeastern Turkmenia, on the Badkhyz upland (600 to 700 m above sea level), steppe cats have been sighted often enough in the semisavannas of extremely hilly plateaus where high ridges alternate with broad open valleys or narrow deep ravines. The sandy loam soil in these places is densely covered with desert sedge and short grasses (myatlik) interspersed with giant umbelliferous fennel (Feru/a). On the crests of high ridges and along their northern slopes the occasional true pistachio tree is encountered. Close to the Gyaz’-Gyadyk mountains this cat is confined to gorges with rocky outcrops, at the bottom of which grow bushes and solitary wild fig trees. In Badkhyz the steppe cat lives in areas where fresh water is not available after April (Heptner, 1956). In the eastern Karakum, near Repetek, this cat is encountered everywhere and confined throughout the year to black saxaul thickets. Occasionally it occupies sand dunes. In summer and autumn the population declines per- ceptibly in the vicinity of Repetek. Possibly some animals migrate in summer to water sources (Sapozhenkov, 1961). In Turkmenia and Uzbekistan this cat regularly hunts in dense reeds and tugais near large water sources and sometimes establishes a den among them. In southwestern Tadzhikistan it avoids dense tugais and is encountered more often in open floodplain forests, especially in winter. It has been met with ‘‘уегу often’’ in the sparse tugais of Tigrovaya Balka (Chernyshev, 1958). Only rarely is this animal reported at sites away from large water sources, and even these are adjacent to springs and wells. Apart from river valleys Photographs of habitats of caracal, sand (saxaul) cat, and in part the jungle cat (tugai) are also a characteristic biotope for the steppe cat. 347 Fig. 201. Тиса! of turanga, tamarisk, and tala willow. Habitat of steppe cat on the Murgab. April, 1962. Photograph by A.A. Sludskii. situated in the plains, in the southern part of the range the cat confines itself to foothills, where it lives in valleys and gorges having sources of fresh water and with arboreal and shrub vegetation. For example, in southwestern Turkmenia the steppe cat lives in foothills and premontane plains but in desert plains occurs in lower numbers. In the sand massifs of Karakum its population is less than that of sand cat. In Turkmenia the cat often is restricted to colonies on sandy stretches not far from rivers, irrigation ditches, or wells. It has also been reported from low mountains (Gyaz’-Gyadyk and Fistashkovyi [ Pistachio] ranges in Badkhyz (southern Turkmenia). It occurs throughout Kopet-Dag at altitudes of up to 2,000 m above sea level (V.G. Heptner). In the mountains of Tadzhikistan and Uzbekistan the steppe cat occurs even to 300 m* above sea level (Chernyshev, 1958; Allayarov, 1963). In Syr-Darya, Chu, Ili, and other valleys of Kazakhstan where this cat is particularly numerous, it is confined to argillaceous desert plains, often covered by solonchaks alternating with tugais, lake and stream systems with reed thickets growing on their banks, and ridges of sandy knolls. Such plains support shrubby tamarisk, Nitraria, salt trees, and rather sparse low reeds. п original text; probably should be 3,000 m—Sci. Ed. 470 Fig. 202. Forest of black saxaul. Biotope of steppe cat in a sandy desert of eastern Karakum. Tolai hare, caracal, and sand cat also occur. Repetek preserve. Photograph by Yu.F. Sapozhenkov. Shrubs grow singly or in clusters, often forming rather impenetrable thickets. Occasionally the areas described above contain individual turanga and oleaster trees, and even small groves. In such areas the lower stratum of vegetation consists of various halophytes, white sagebrush, grasses and forbs. In some years these areas are inhabited by large numbers of tamarisk and midday gerbils, domestic mice, tolai hares, and pheasants. Apart from argillaceous sections of river valleys cats are often encountered in small tugais and among reeds by water sources to which they repair in order to hunt. At such times they occasionally may be seen totally exposed on banks of rivers and streams. Characteristically, in sand knolls adjoining river valleys, and more so in extensive sandy massifs in Pri-Balkhash (for example, in the Taukum and Sary-Ishikotrau sands), the spotted cat is absent or encountered rarely, and chiefly in winter. In winter one may encounter it among saxauls 10 to 20 km away from rivers. In such places where saline plains often alternate with sandy ridges this cat distinctly prefers the former and is rarely seen in the sands. It avoids completely exposed places and hence tracks usually run through sparse thickets or along their edges. When living in thickets, the 348 471 cat while hunting regularly follows trails of roe deer, wild boar, hare, and pheasant. In summer, if living close to water sources, it is not afraid of crossing them as it is an excellent swimmer. Cats have been seen on several occasions, without visible cause, swimming across the Ш River (several hundred meters), and its tributaries, which have rapid currents (A.A. Sludskii). In the foothills of Talassk Alatau this predator lives in gorges with a vegetative floor of willow, sea buckthorn, and poplar, or tugais of oleaster and elm with an undergrowth of dog rose, barberry, honeysuckle, and blackberry. Rarely, it is met with in forests of wild apples with an understory of bushes. In winter the steppe cat hunts in gorges and on sunlit rock ledges and steppe slopes where mouse-like rodents, and rock, bearded, and gray partridges are common. In these mountains it usually ascends to 1,300 т above sea level, rarely higher; in winter, however, it is confined to much lower levels. For example, in Aksu-Dzhebaglinsk preserve a steppe cat was once caught in summer at а height of 1,800 т (Т.А. Yanushko). In the foothills on the northern slopes of the Kirgiz range this cat inhabits the lower parts of gorges densely grown with dog rose and other shrubs. In the Chu-lli hills, where there is depauperate grassy vegetation of a desert or steppe type, the animal lives in gorges along the bottom of which flow streams densely overgrown with vegetation. It ascends there up to 1,500 m above sea level. Under the even more severe conditions prevailing in the Dzhamby! mountains, steppe cats live on slopes which during summer are green but support only scattered bushes of boyalycha, sagebrush and sheep’s fescue. In this region only in the gorges are springs sometimes found, which usually dry up by autumn; along the banks of these springs tamarisk, salt trees, and cheegrass [Lasiagrostis| grow. Great gerbils [Rhombomys] shelter in these gorges. In the foothills of the Trans-Ili and Dzhungarsk Alatau the cat lives along river valleys with tugais or willow thickets; it is rarely encountered on slopes covered with desert or steppe vegetation, or in rocks. In the mountains it does not, as a rule, ascend beyond 2,000 m above sea level. For example, in the Chilik valley in the Bartogai region, situated at a height of 1,600 m above sea level, steppe cats live in dense tugais of turanga, oleaster, sea buckthorn, willow, and reeds, among which are scattered extensive openings covered with dense grasses and forbs. This region also contains many tolai hares, pheasants, and mouse-like rodents. In Ustyurt and Mangyshlak the cat lives throughout the year along rocky or argillaceous precipices, in chinks [arroyos] close to sources of fresh water and overgrown with tall herbage, reeds, and scattered bushes, on sand knoll islands, and saxaul thickets, and often deep inside the desert 80 to 100 km from chinks and the nearest fresh-water source. In Kirgizia the cat is common among reeds and sea buckthorn thickets 349 Fig. 203. Slopes of northern Chink and Er-oilan-duz depression. Habitat of steppe cat in Badkhyz desert. Southern Turkmenia. February, 1961. Photograph by M.V. Heptner. in the floodplains of the Chu and in Issyk-Kul’ depression (D. Dement’ev, 1940; Zimina, 1953). In the western part of this depression, at Rybachii village, the steppe cat lives in cheegrass stands (Lasiagrostis), often growing to a man’s height, and is common on the northern shore [of Issyk-Kul’] among sea buckthorn shrubs (A.A. Sludskii). In the Talas valley it lives in thorny shrubs (salt trees and dog rose) or in poplar forests in the river basin (Kuznetsov, 1948). On the slopes of the Kirgiz range it is found mainly in the lower montane zone in dog rose thickets, but sometimes quite high in the mountains. A kitten of this species was caught in the upper Karabalta at a height of about 2,500 m above sea level (D. Dement’ev, 1940). In the Mongolian People’s Republic the most typical biotopes of this cat are sandy knolls in depressions overgrown with dog rose and saxaul shrubs in sandy soils. Ц does not ascend to high altitudes in the mountains (Bannikov, 1954). Throughout the range within the Soviet Union this carnivore does not avoid human settlements, and often colonizes oases, or settles in gardens on the outskirts or close to settlements, especially in winter. It is attracted to such places by water and mouse-like rodents and often by pheasants. 473 Fig. 204. Habitat of steppe cat in Badkhyz desert. Meadow slopes densely popu- lated by great gerbil (Rhombomys opimus). Southem Turkmenia. February, 1961. Photograph by M.V. Heptner. The cat, as mentioned above, prefers to live close to fresh-water sources, but at some places lives year-round even in waterless deserts; the absence of natural fresh-water sources does not restrict its distribution. However it is poorly adapted to living in regions with deep, soft snow cover. The weight load per cm? of resting surface on its foot is equal to 90 to 120 g and the height of the forelegs from the ground to the chest is 18 to 23 cm. Hence it can only negotiate soft snow cover 20 cm high or more with great difficulty, dragging its abdomen in the snow. Moreover, it cannot get at rodents from under the snow. In the lower reaches of the Ili during fresh snow these cats were seen taking advantage of hare trails leading to their forms; on the second day after the snowfall this animal had made numerous trails and its predator used them regularly. Thus the presence of a large number of tolai hares in the floodplains of desert rivers makes it possible for this cat to move easily through deep, soft snow cover. In the foothills the predator avoids deep snow and confines itself to sunny sections and rocky precipitous slopes of gorges (A.A. Sludskii). If a snow cover 20 to 30 cm high or more persists over a long period, it acts as an important 474 JEAN OTT 0`0Е (5 риЯ РТ — — — [snjiano snulysaiuaz]] Зоцэзрэч poivs-3u0'] lez — — — dooys [nyeiey jo заше] a OTT O's OT рэупиэр! 1э43113 jou ‘sjuspoy 176 — — — [snupy3fo ‘И эюл ичеч8 ту Vel =: = =- [5714013] 2104 so sz — Cz — э5зпош э5поН — — os — [sno1upjsayany `У] тел ива, ve ae TE Tol [iqio3 Aeppryy [6 0`РТ SLI ae [Iq193_pa[ley-poy [с 0'C7 ae TBE 118 3910) C9C СТР CLI == $19190 gs 8 Е (as sevoqiof = Е 0'0€ СР 9116$ рипо18 рэоз-шч1, orl = CL 9°6 этеЧ тео, 9`96 — SLO — ззиэрол рие $Ч4лошо8е] == 019 SLO = ее sojdues cpt sojdues 9¢ $э1Аше$ Op sojdues 6 (9961 ‘охизлеа (L961 (861 (1961 рие urunysy) ‘aoleyysey ‘@) ‘лэцзАилэцо) ‘aoyuoyzodes) yajoday wnyAzAy и1215$еЯ э44э1$ ysurysiey unyeyxysey ‘TUNyeIVy Woyseq 1851292] “URISTIYZPe т, sojdues Jo 1эашпи ‘uoIsay (sojdwies Jo 1aquinu [е30} Jo эЗезизэлэЯ se pajuasaid sadaq PUB $}4э}и0э YDLUIO}S JO SISA[BUR) BISY эр! JO Sjlesep WisyyNos Ul ye eddas Jo роол “ST эЕТ, Tse 475 ТТ "ря 'ю$—вэр jou Зишеэи, “PA `125$ — рле2 | patynuapiuy) „ — $э^еэ| 55818 иээло) == +*5р99$ 85 — зитешол JULI g GE зрузивеча — $12429 © — sioddoyssein 96 sI9yjO рие $э1ээа yorl|g — рэчуциэрг 143113 jou ‘sayo0q 8<Т spodomyyiy — $339 эзюло, ее [opnjsa J] 2810}10) 11959 СЕ [6424110 ‘aydozq] эхеиз зе — [snjojoauy siydowwosg] ayeus Moly ЕС рэцпиэр! 12431} jou ‘sprezry] [514025024 2:6 5п]руаэоои“ац | зреэцрипо1 Aye J, — ,DyAnyoysvd Pour] OT [pjuajoumnsuvs ошь8у] ewese э44э16 == dasys PTI = $2139э си $882 риа — sjuesvoyd OT [21210540 sky2adopoyy] youy yesoq СФ [27215142 0р142]00] Уле| рэ1$э1о ЕЙ $91955 4 350 476 factor limiting the distribution of the steppe cat (Formozov, 1946), leading to its absence in zones of semideserts and steppes in the plains and in the upper montane zone. The steppe cat is quite sensitive to low temperatures. Three cats held in a large cage with hay at ап air temperature of —30°С for one night had frostbitten ears and one even a partially frostbitten tail. Cats which fall into traps suffer frostbite (A.A. Sludskii). In spite of this the steppe cat inhabits regions with a sharply continental climate. In the northern parts of its range the winter temperature sometimes falls to -35° or -40°C. The anima