UU.A..A. SSEMENOVEMENOV

UMAR AUBEKIROVICH SEMENOV

Candidate of Biological Sciences, author of more than 40 scientific papers and several monographs, including “Biological Monitoring in Karachay- Cherkessia”, “Wild Boar in the Teberda Nature Reserve”, “West Caucasian Tur in the Teberda Nature Reserve”. U. A. Semenov has developed and implemented the method of organizing and conducting systemic biological monitoring on protected natural areas. Since 2010, U. A. Semenov has been the head of the Persian Leopard Breeding TTHEHE WWISENTSISENTS and Rehabilitation Centre of the Sochi National Park Federal Governmental Budgetary Institution. Research interests: ecology and ethology of the ungulates and predators of the Caucasus. OOFF KKARACHAY-CHERKESSIAARACHAY-CHERKESSIA E-mail: [email protected]

Photo by R. Tekeev U.A. SEMENOV

THE WISENTS OF KARACHAY-CHERKESSIA

Proceedings of the Sochi National Park Issue 8

KMK Scientifi c Press Moscow ™2014 UDC 599.735.51(470.63)

Semenov U.A. The Wisents of Karachay-Cherkessia. Proceedings of the Sochi National Park. Issue 8. Moscow: KMK Scientifi c Press. 2014. 171 p. This study represents the history of wisent reintroduction (Bison bonasus L.) in the Teberda State Nature Biosphere Reserve, its historical expansion and occurrence in Karachay-Cherkessia between the 18th and the 20th centuries. The book describes and analyses the population dynamics, changes in the migration patterns and social structure of the species in the mountain environmental system. The author has summarized the records of the Teberda Nature Biosphere Reserve since 1968 and shown the results of the fi eld observations: diurnal activity, spread within the habitats, diet, reasons for extinction, wisent behaviour in the course of adaptation and further exploitation of the upper part of the Kizgych Valley of Arkhyz. The book is advisable for specialists in the area of protection and management of wild ungulate populations, as well as for other readers interested in different environmental, ethological and biological issues related to wisents.

Science Editor: B.S. Tuniev, Merited Environmental Specialist of the Russian Federation, Doctor of Biological Sciences

Reviewers: T.P. Sipko, Head of the Wisent, Bison and Muskox Research Committee in Russia of the Russian Mammalogist Society of the Russian Academy of Sciences, Senior Researcher of A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences, Candidate of Biological Sciences N.V. Treboganova, Head of the Wisent Nursery of the Prioksko-Terrasniy State Nature Biosphere Reserve, Candidate of Biological Sciences

The study is published upon the decision of the Council for Science and Engineering of Sochi National Park Federal Governmental Budgetary Institution, Minutes No. 6, dated June 6, 2013

This publication was supported by WWF-Russia

© U.A. Semenov, text, illustrations, 2014 © Sochi National Park, 2014 ISBN 978-5-87317-984-8 © KMK Scientifi c Press, 2014 In loving memory of my father

Introduction The problem of wisent reintroduction in Karachay-Cherkessia remains acute even 44 years upon their release in the Arkhyz area of the Teberda Nature Biosphere Re- serve; moreover, it has become tragic. In the 1990s, extinction threatened local popula- tions of the Caucasian region again. The economic and political instability in Russia nearly whittled down a splendid example of sophisticated and coherent policy of popu- lation recovery within its historical habitat, as well as monumental efforts of several generations of scientists and nature reserve workers. The wisents as very large animals became easy prey for poachers. It resulted in a critical drop of the wisent population in some areas and in their complete extinction in the other areas. The fate of the wisent population of Karachay-Cherkessia was not different… The wisent as a species of the archaic epoch has managed to survive until our days and is extraordinarily valuable as an element of the fauna diversity of the Karachay- Cherkess Republic. The wisent came into the spiritual life of local people as an element of tales of the powerful natural forces many centuries ago, and now it remains a na- tional treasure. Its example contributes into development of the environmental culture of the modern society and national nature protection policy. Most unfortunately, the native wisent extinguished from the Caucasian fauna for- ever, but consolidated efforts of the global nature protection society of the 20th century allowed for the species preservation for the future generations. Therefore, the animals are being returned to their initial habitats, including Karachay-Cherkessia. This publication is focused on the analysis of the wisent reintroduction history in the Kizgych Valley of Arkhyz since their release and until 2012, inclusively. Various environmental and biological issues are analysed on the base of the Nature Records of the Teberda State Nature Biosphere Reserve from 1968 to the present day, archive records, scientifi c studies, and own observations between 2000 and 2012. The space- time analysis of the population dynamics allows for objective assessment of the popu- lation development sustainability within 44 years by means of defi nition of the most important factors infl uencing the wisents in the mountain environmental systems of the North-Western Caucasus. The data are arranged chronologically in order to analyse the animals’ adaptation to the new conditions and understand the problems they faced af- terwards. In studying such issues as population dynamics, migration habits within the nature reserve and its surroundings, social structure, habitat location, reasons for ex- tinction, etc., we also tried to fi nd out biotic, abiotic and anthropogenic factors mostly infl uencing the Arkhyz wisent population inhabiting the highlands that are deemed most complicated for living. 6 U.A. Semenov The Wisents of Karachay-Cherkessia Our country has been dealing with the wisent population reintroduction for more than 60 years now: special nurseries have been built, and programs and strategies of wisent release to new areas have been developed. Still, the wisent (Bison bonasus bonasus Linnaeus, 1758) as an endangered fauna species remains on the IUCN Red List of the Russian Federation. Now Russia has different economic conditions and principles of interrelation between the Federation and its subjects, as well as access to the resources of the other countries; thus, the wisent preservation strategy in Karachay- Cherkessia shall be developed with regard to the species reintroduction analysis of the Teberda Nature Biosphere Reserve. The Arkhyz population was unable of independent reproduction and development in 2012. The fi ve remaining wisents were living their last years, and Karachay-Cher- kessia could have lost again a most important element of its fauna diversity. However, upon several complicated discussions of the new wisent preservation strategy in Russia (including the expediency to release new species in Arkhyz), eight new wisents were again delivered to the Kizgych Valley from the Okskiy State Nature Biosphere Reserve. It was the achievement of the supporters of the Kizgych group recovery. Herewith the initiative put forward by WWF in Russia and supported by the Ministry of Natural Resources and Environment of the Russian Federation was of great importance. Therefore, the release of new animals became a new milestone in the wisent history of Karachay-Cherkessia. It also gave us a unique opportunity to carry out comparative analysis of the wisent adaptation and their further life in the new environmental condi- tions. We hope that it will be the fi rst step of a long-term program, the development and implementation of which is required for sustainable development and qualitative management of the Arkhyz wisent population. Now that the book is fi nished the author has the honour to thank everyone who has helped him in the course of writing this study. I would like to sincerely thank the family of Madzhir Lokmanovich Khubiev who had worked for the nature reserve for more than 30 years and passed untimely. His principles that sometimes used to expose him to the direst dangers have played the key part in confrontation with poachers and illegal deforestation. I would like to thank his son Ramazan Khubiev who continued the famous traditions of protection of the unique Kizgych Valley as a forester and did his best to help me working in the mountains. His professionalism, understanding of the wisent reintroduction importance and sincere desire to preserve wisents contributed to making a positive decision on the wisent re- lease in Arkhyz in 2012. I am very grateful to Seitbiy Akhmatovich Dzhanibekov, a state inspector of the Kizgych Forestry, as well as my guide and companion, who has spent so many hours and days on this research so far from his family and home. His incredible anxiety for knowledge and unconditional love for the mountains helped me so much in compli- cated expeditions and collection of data on the environmental and biological issues of the Kizgych Chine. Introduction 7 The data given to me by Grigory Yakovlevich Bobyr, who had worked in the moun- tains for a long time and dedicated his life to research of the Karachay-Cherkessian fauna, helped me a lot in writing the study. Our fi eld observations played a signifi cant part in understanding several environmental and biological aspects of wild animals in the course of cooperation at fi rst in the Teberda Nature Biosphere Reserve and then in the Sochi National Park. I would also like to thank Nikolay Dmitrievich Penkovskiy, the Director of the Sochi National Park, who has always supported me in successful researches and publication of this study. I also am sincerely grateful to B.S. Tuniev, N.V. Treboganova, E. Tsibizova, and T.P. Sipko, who agreed to edit the study and gave me most valuable remarks. I deeply appreciate the monumental patience of my family and their efforts to create the conditions required for my long-term stay in the mountains and my favourite job!

Umar Semenov Chapter 1. Evolution and Distribution of the Genus Bison H. Smith, 1827

The European bison (wisent) is the heaviest surviving bovid among the wild ungulates living in the Caucasus now. Nowadays species of the Bison genus live in different conditions: from woodland valleys to mountain forests. Un- fortunately, there are a lot of gaps in the studies of the European and American bison related to their wild life. Although there are many publications on ecology and biology of these animals, they are more or less dedicated to separate parts of their former habitat and can hardly refl ect all the variety of life cycles and adaptations of these animals inherent in them 100–200 years ago. For example, no large-scale migrations of the bison in the American prairies happen nowadays the way it used to be before Europeans came to America. Despite of the fact that many characteristics of the behavior, ecology and biology of the species have been preserved, they have changed signifi cantly because of the decrease in the biotopes’ size and variety, change in the qualitative composition of the feed, time of the rutting season and birth of calves, etc. Nevertheless, the wisent as a living ancient monument of nature which has survived until present has always attracted the scientists with its noble strength. Thorough analysis of the fossils and the information in the literature has made possible for a group of scientists to restore many features of the external appear- ance and way of life of the European and American bison inherent in them in the past. The complex monograph (Wisent, 1979) provides a systematic description of the modern and extinct bison (Bison) of the Late Pliocene (approximately, 2 mln years ago) and the Early Pleistocene; the latters have distant resemblance with the present day species. They were small and primitive bison (typical repre- sentative: Bison palaeosinensis Chardin et Piveteau, 1930), which, presumably, dwelled in the warm climate forest area, fed on woody vegetation and forest grasses with soft epidermis. Later in the Pleistocene epoch from about 1.6 mil- lion to 10–11 thousand years ago, there were animals of different sizes within the species: from small to big ones. A signifi cant difference occurred due to the animals’ feed type, because of which they developed relatively big teeth. In the Early Pleistocene and Middle Pleistocene periods there were European and Asian forest bison of medium and relatively small sizes, as well as a sig- Chapter 1. Evolution and Distribution of the Genus Bison 9 nifi cant number of bigger animals (Bison (Bison) voigtstedtensis Fischer; Bison (Bison) schoetensacki Freudenberg). The researchers have not discovered the complete morphology of Bison schoetensacki, it is only known that its bones are thinner and longer than those of other species of the Bison genus. Presum- ably, they were widely spread in Europe (except for its northern part) and in the temperate zone of Asia (bone fossils of this species were found in the East up to the Minusinsk Hollow and the Vilyuy River basin), the species might have dwelled also in the north-east of Yakutia. With that the Siberian rogues of Bison schoetensacki were bigger than their European congeners. It is possible that the Asian population further on formed a separate geographic race. The bison of the end of the Middle Pleistocene and Late Pleistocene and the modern Asia-Canadian branch are represented by several species, the most typi- cal of which is Bison priscus Bojanus. The exception from this race is Bison priscus gigas, which occupied mainly the southern border lands of the distri- bution area of Bison priscus in the steppe zone of Asia and the eastern part of Europe. The external appearance of late representatives of Bison priscus was almost completely restored in the middle of the 20th century, due to many fi nd- ings of whole bones and even complete skeletons and corpses in the perma-

Figure 1. Late Pleistocene Bison (Bison priscus occidentalis) (after Flerov, 1979). 10 U.A. Semenov The Wisents of Karachay-Cherkessia frost zones of the Eastern Siberia and Alaska. Within a historically huge habitat of Bison priscus, separate parts of the population developed (sometimes for a long period) under the infl uence of certain environmental factors peculiar only to those regions. As a result of that process new ecological races appeared, and quite often they were signifi cantly different from each other. It is enough to say that there were forest, forest-tundra, forest-steppe and even steppe forms that preconditioned peculiar feeding adjustments and corresponding morphological adaptations. From the end of the Mindel-Riss (about 150 thousand years ago) to the be- ginning of the Würm (about 70 thousand years ago), bigger bison (Bison pris- cus crassicornis) lived in the Northern part of East Asia, in the west up to the Yenisei, and also in Alaska and Canada. This species had a higher level of adaptation to the severe conditions of forest-tundra, forest-steppe and cold crook-stem forest. A range of characters (for example, great width and volume of the nasal cavity, etc.) proves that those animals were adapted to live in cold conditions. In the opinion of the researchers (Flerov, 1979), the exterior parameters, which they have managed to defi ne based on the skeleton dis- covered in the permafrost strata, are determined by the following characters: high-lifted head, a very tall distinct hump, but the back is almost parallel to the ground and does not have a fl at bend towards the hind quarter. Taking into account that the main habitats of the animals were cold regions, their coat was characterized by thicker and longer fur. The largest of all the European and Asian bison (tip-to-tip spread of the horns reached 2 meters and more) of that epoch lived in the southern part of Eastern Europe, Kazakhstan, Southern Siberia and Northern Mongolia. There are de- scriptions of quite many forms of this species close in size and living in the simi- lar landscape and climatic conditions. These are Bison priscus latifrons and Bi- son priscus gigas living in wide and open spaces. The researchers found only the skull without the lower part of the latter but the analysis of the glenoid surface and the simulation of bone articulation let them conclude that the animals had been adapted to eat hard steppe grasses and as a consequence bound to steppes of warm or temperate climate. Fragments of rock paintings of animals played a great role in the understand- ing of the bison’s evolution. Some of them depict the external appearance of those animals quite truly. Some of the most impressive paintings created by the prehistoric human being are the images of the bison discovered in the famous cave of Altamira in Spain, Font-de-Gaume, Les Combarelles and Lascaux caves in France, which pertain to the period of the Aurignacian, Gravettian, Solutre- an and Magdalenian cultures dated from 35,000 to 8,000 years B.C. The fi rst features in the rock paintings corresponding to the morphological characters of Bison bonasus appeared within the Magdalenian period dated from 12,000 to Chapter 1. Evolution and Distribution of the Genus Bison 11 8,000 years B.C. According to the authors (Flerov, 1979), as compared to the bison, they have more uniform colouration, a smaller beard, a less developed hump and a mane above the neck and on the crest. In accordance with the opin- ion of the scientists, the Late Pleistocene bison were very similar to the bison of the Eastern Siberia and North America of the same time. Thus, successive historical epochs characterized by changes of environmental factors, which in their turn contributed to the development of various adaptations in the process of evolution, generated various bison characters which helped them to overcome hardships, favoured differentiation based on the type of diet, and developed preferences in the biotope location. Different parts of a huge his- torical bison habitat stood in marked contrast with each other in terms of their landscape and climatic conditions that brought about the formation of several local races. It was assumed that possibly only in the territory of Canada and the USA there were several local races: plain race in Kansas and North Montana; forest race in the eastern part of the habitat within the forest area, and mountain race in the western mountain part of the species habitat. Remains of the European-Caucasian branch of Bison, ancestors of the pres- ent day wisents (Bison bonasus), were found almost all over Europe (except for its northern part), the Greater Caucasus, the South Caucasus and in the Northern Iran. In the Caucasus region the most ancient remains of small bi- son were found in the Upper Pliocene Absheron clay of Kabristan and in the strata of Taman Peninsula (Burchak-Abramovich, 1951, 1957; Vereshchagin, 1957, 1959). Therefore, the Caucasian wisent (Bison bonasus caucasicus) was the direct descendant of the local Pliocene-Pleistocene Bison. According to N. Vereshchagin (1959), Bison’s degradation in size is mistakenly thought to be related to the climate change and temperature decrease in the Upper Pleisto- cene, whereas the temperature decrease, quite the opposite, had conduced to the increase of the wisents’ size in the Middle Pleistocene. The wisents’ size decrease was caused by the post-glacial fragmentation of the population and concentration of animals along river valleys, in the highlands and broad-leaved forests of the mountain slopes. Isolated development of the Caucasian part of the wisent population and their gradual adaptation to mountain conditions led to signifi cant differences in the morphology and life habits of the modern wisents as compared to their ancestors living in wide open spaces and dry steppes of Eurasia. For example, many bison living in the North American prairies were capable of running fast and for a long time, they were enduring and strong. A horse rider could not run down a herd of running bison. But the wisents living in the forest contrariwise could not run fast and long, as the con- ditions (dense forests and wind-broken trees) did not favour the development of those capabilities, but the wisents could jump high, and that was essential for the animals in such habitats. 12 U.A. Semenov The Wisents of Karachay-Cherkessia Until the Middle Ages the wisents’ habitat probably also included all the areas of the Caucasus Isthmus and the eastern part of Europe along the Don Valley, but fast-growing factor of pursuit by humans and extermination of the original habitats of the animals led to complete extinction of those animals in the easy- to-reach steppes and forest-steppes of Eastern Europe and Asia. Huge armies of the Khazars, the Cumans, the Mongols, etc. easily trapped and killed such large animals as wisents during massive driven hunts within large territories. Comparative analysis of limbs of the bison living in the open spaces and the wisents living in the forests is pretty interesting. It is known that the limbs of the latter are longer, but at the same time short-legged bison are capable of running fast and long while wisents are not. According to the results of comparative anal- ysis, forest-dwelling animals almost always have longer limbs than their fast- running congeners living in the open spaces. For example, the red deer as a typi- cal forest-dweller has longer legs than the tundra-dwelling reindeer capable of running fast and long. According to the scientists (Wisent, 1979), this peculiarity of adaptation of different forms lies in the position of the center of gravity. The longer the limbs are, the taller the animal is, and the bigger the vertical lift com- ponent is needed for pushing during running. Probably the constant necessity to stretch forth while looking for a support between the stones, logs or soft snow on the slopes played an important role in lengthening of the wisents’ limbs. In the plain forests the animals live in different conditions, and it has shown through their evolutional adaptation to move constantly with minimal vertical variations.

Figure 2. North American Steppe Bison “Barmen” Aged 9 Years, in the Prioksko-Terrasniy Nature Reserve. Photo by N. Treboganova. Chapter 1. Evolution and Distribution of the Genus Bison 13 This specifi c feature of mountain animal forms, including the Caucasian wisent, formed a signifi cant difference in gracefulness between the Caucasian mountain wisent and the European forest wisent with the course of time. Ancestors of the modern wisents, ancient primeval representatives of Bison genus, mainly occupying plain forests, forest steppes and even steppes, once had a wide range. They were found from the shores of the northern seas to Europe, were common in North America, they inhabited India, China, south of Siberia, Kazakhstan (Kulagin, 1918, 1919; Gromova, 1935; Vereshchagin, 1957, 1959; Heptner et al., 1961; Sokolov, 1964; Burchak-Abramovich, Vekua, 1980, 1999, and others). The huge geographical range of the Bison genus was formed in the process of evolution and changes of the animals’ living environment. In some places they appeared adapting to the new environmental conditions, in the other places they disappeared because they could not survive in such changed condi- tions. In the Low Pleistocene early forms of bison occupied the whole Siberia, then they were replaced by the new, more adapted forms which occupied almost the whole Palearctic. At the same time some part of them disappeared from the territory of the Southern and Southeast Asia, and eastern populations entered North America and south of Eurasia via the Bering Land Bridge in the Middle Pleistocene (Flerov, Zablotskiy, 1961). The summary report of A.A. Danilkin (2005) provides information about historical habitat of the ancestors of the present day wisents in the territory of Russia. The remains of primitive bison were found in the Pleistocene stratum

Figure 3. Bull of the Lowland-Caucasian Bison (B. b. bonasus + B. b. caucasicus) “Shponty” Aged 15 Years, in the Prioksko-Terrasniy Nature Reserve. Photo by N. Treboganova. 14 U.A. Semenov The Wisents of Karachay-Cherkessia of Priazovye, the Crimea, the North Caucasus, and the regions of the Krasno- dar Territory, Rostov, Astrakhan, Orenburg, Volgograd, Saratov, Samara, Penza, Ulyanovsk, Voronezh, Orel, Kursk, Bryansk, Moscow, Ryazan, Tver, Vladimir, Leningrad, Novgorod, Pskov, Vologda, Arkhangelsk, Kirov and Perm, as well as the Republics of Komi, Tatarstan, Bashkortostan, Mordovia, some areas of the Middle and Southern Ural, Trans-Urals, East and West Siberia, South Primorye. Fossils were also discovered in Moldavia, Ukraine, Belarus, Kazakhstan, Kyr- gyzstan, Uzbekistan, and in all the republics of the South Caucasus. Early ances- tors of the modern bison and wisents were signifi cantly smaller and had distant resemblance with the modern ones, but in the process of evolution their external appearance has been constantly changing developing certain adaptations. In the later cold period when the animals inhabited wide open spaces of the Northern Eurasia they were large and even huge but in the course of time they became smaller in size. Cold weather recession brought about growth of new greenery and change of landscapes, and along with that even the animals belonging to one and the same species but living in the different parts of the huge habitats changed in different ways with the course of time because they were infl uenced by various combinations of environmental conditions. Authentic remains of wi- sents themselves started to appear only in the Holocene in Europe, including the Caucasus, and presumably in Trans-Urals and West Siberia (Danilkin, 2005). Many researchers (Gromova, 1935; Flerov, 1955; Sher, 1971; Rautian, Sipko, 1997; Rautian et al., 2003, and others) think that from the point of view of the habitat type and peculiarities of the diet, the genus (Bison) may be nominally di- vided by the environmental specialization type into the forms inhabiting forests (also including mountain forests) and those inhabiting steppes. The forest-dwell- ing forms were further on defi ned as wisents, and the steppe-dwelling forms were called bison. Due to the differences in the diet, animals of the fi rst and the second branches developed certain adaptations (structure of intermaxillary bones, teeth structure and external morphological characters), however, as rea- sonably stated by A.A. Danilkin (2005), adaptations to different types of habitats are not absolute and often have only adaptive nature developed by the animals in the course of evolution. Nevertheless, the issue regarding the taxonomy of one or another form is still open. Defi ning the differences based on a range of characters, the scientists (Flerov, 1965; Heptner et al., 1961; Bohlken, 1967; Sokolov, 1979; Krumbiegel, 1980; Groves, 1981) separate three species, inside of which they also name two sub-species of bison, differentiating them as a steppe bison and a wood bison. However, modern molecular genetic analyses did not reveal any signifi cant dif- ferences. Comparative analysis of the wisent and the bison lets us determine a number of distinctive characters in morphology, colouration, skeletal muscles and skull Chapter 1. Evolution and Distribution of the Genus Bison 15 structure. However, the results of biochemical, cytogenetic and molecular ge- netic tests have shown that the bison and wisent are almost identical. Compara- tive analysis has shown that behavior, social structure and general environmental and population-related characteristics are similar in many aspects. Longstanding work to restitute wisent which has been carried out in many countries has shown that the animals interbreed and give birth to the calves capable of reproduction. Though there is no agreement among taxonomists regarding the taxonomic sta- tus and composition of the genus (Bison); fi nding the answer to this question is one of the main tasks for defi ning the further strategy of the wisents’ preserva- tion in the wild. Chapter 2. The Wisent in the Caucasus

2.1. History of Studies of the Caucasian Fauna

The fi rst targeted zoological researches of the Caucasus were started in 1770 by academician Anton Güldenstadt (Güldenstadt, 1787), the result of his travels was the description of new animal species (including the West Caucasian tur — Capra caucasica). A bit later special expeditions led by Samuel Gmelin, a member of the Academy of Sciences, and Marschall Bieberstein worked there, and in 1793 the Caucasian region was visited by Peter Pallas who summarized the collected infor- mation in his “Zoographia Rosso-Asiatica” in 1811. Only since the 1720s different zoologists started to visit the Caucasus: E. Eichwald, E. Menetries, E. Eversmann, A. Nordmann and others. Being a totally non-explored land, the Caucasus attracted many natural scientists of that time. However, the possibility to perform detailed zoological research was signifi - cantly limited by the complicated political situation, diffi culty to approach and baffl ing complexity of working in extreme conditions in the remote mountain chines with a danger to life, and that left a mark distinctive for many publications of that time. Later on zoological researches were made by the Caucasian Mu- seum or by the Caucasian Department of the Imperial Russian Geographical So- ciety. Regular expeditions to different parts of the Caucasus led by such talented scientists as N.A. Severtsov, K.A. Satunin, A.N. Kaznakov, A.B. Shelkovnikov and others signifi cantly enriched the science with new knowledge about species composition of the animals, certain special features of their geographical expan- sion and mode of life. “The fi rst explorers of the Caucasus, as it always happens in such cases, tried to travel around as large area as possible, visit as many interesting places as possible, amass collections and enrich the science with new species of animals and plants, but of course they could not study in detail the peculiarities of the Caucasian fl ora and fauna, study the varieties and make a detailed comparison of the Caucasian animals and plants with their relatives inhabiting the neighbouring countries of Europe and Asia, and study their geographical expansion within the Caucasus and the neighbouring countries, i.e. they could not engage in the studies which were very important for clarifi cation of many interesting issues of general nature. That is Chapter 2. The Wisent in the Caucasus 17 why based on the studies of the fi rst explorers of the Caucasus it was only possible to conclude that the Caucasian nature was very rich and diverse; and only funda- mental research of the last 20–25 years has shown that studying the fl ora and fauna of this country, which was crossed not only by the routes of migrating peoples but also by the routes through which different animal species spread occupying wider areas, is still very interesting from the scientifi c point of view, throwing light on the distant past of the modern Europe and Asia, on how and from where the Cau- casus has been inhabited by the plants and animals, on what the routes of animals’ displacement in the current conditions and during previous geological epochs have been, on why the fauna of the known part of the Caucasus has this and not some other face, etc.” (Dinnik, 1911, p. 2). We can probably consider that the fi rst survey of the Caucasian fauna was a tractate by E. Menetries dated 1832 called “Taxonomic Catalogue of Zoologi- cal Objects Collected during Travels around the Caucasus”, containing a perfect summary of all the information known at that time about the Caucasian wildlife. According to K.A. Satunin (1903), this tractate has been the only research for 67 years that has covered all fauna of the Caucasus. Many natural scientists basing on their own research tried to trace the ways of the Caucasian fauna formation during the historical epochs, and it very often led to confusion because of the main challenge: limited knowledge and little in- formation from different parts of the Greater Caucasus. One of the fi rst attempts to divide the Caucasus into zoogeographical areas was made by K.A. Satunin (1903) in his article called “Review of the Studies of the Caucasian Mammals”, which was published in the January volume of “Publications of the Caucasian Department of the Imperial Russian Society”, and in 1910 at the general meet- ing of the Department members he made a preliminary statement where he in- formed that phytogeographical regions determined by botanist A.S. Medvedev in 1907 coincided with the zoogeographical areas which he had defi ned. In his study “Ideas of the Caucasian Fauna Origin” K.A. Satunin wrote: “It is clear from these two studies that the Caucasian region is not something integral from the point of view of both fl ora and fauna, it is a combination of different fl oras and faunas of absolutely different origin. That was the cause for that mess and discordance which were found in the studies of all the zoogeographers when they tried to force the whole Caucasus into one of the parts of the Palearctic region which they accepted. It became absolutely obvious that the phrase “fauna of the Caucasus” was as uncertain as the phrase “fauna of the Russian Empire”. Unfortunately, many people who are not well-informed still use the word “the Caucasus” in the sense of some integral zoogeographical unit, and of course that leads only to a mess” (Satunin, 1910, p. 29). K.A. Satunin visited Karachay-Cherkessia in 1913, at that time his son and he crossed the Sanchar Pass from the headstreams of the Bzyb River (Abkhazia) to 18 U.A. Semenov The Wisents of Karachay-Cherkessia the Zagedan Valley (headstreams of the Bolshaya Laba River), then he travelled to the headstreams of the Kuban River where he explored such isolated terrains as Uzun-Kol, Dzhalan-Kol, Daut Chine and others (Satunin, 1916). Separate parts of the Greater Caucasus have different origin and age. Geologi- cal structure of the region clearly shows the elevation of the Earth crust strata in different epochs. The fauna of the region formed in correspondence to that. As the changes were happening within a long time the animals could adapt to that territory. The most ancient part, the Central part of the region, the Greater Caucasus Mountain Range and its spurs preconditioned the presence of the great amount of endemic species of plants and animals in the Alpine zone while the forests and lowlands were often inhabited by the incoming species much later.

2.2. Studying of Wisents in the Caucasus

In the Caucasian region the wisent has always been considered a native spe- cies. Bone fragments of the ancient wisents (Bison priscus) were found in many parts of the Caucasus through to Talysh and the borders of West Asia. The earli- est written records on wisents’ life there are found in the ancient tractates dated the beginning of the 13th century describing hunts of Abaqa Khan in Aran during the winter of 1275/1276, and Ghazan Khan in Talysh in 1301/1302. At that time Rashid al-Din, the Iranian chronographer of the 14th century, de- scribed massive hunts of Ghazan Khan, “Emperor of the Islam”, who was one of the Mongolian rulers of Iran in Aladag, and mentioned that the khan who had stopped in the quarters in the South of Kura-Aras Depression and Hamshahri was hunting in the mountains during his way to Talysh and Ispakhbad. “Ghazan Khan ordered to build in those mountains two wedgewise converg- ing hedges made of poles and brushwood so that it would take one day to walk along them, so that it would take about one day to cover the distance between the ends of those hedges in the wide part of the wedge, and so that the distance between them in the narrow part would be fi fty gyazes, and to make out of a tree some kind of a paddock in the hedge blind end. After that the soldiers organized stalking and wedged mountain buffalos (here he means wisents (Nemtsev et al., 2003)), djurs, wild goats and donkeys, jackals, foxes, wolves, bears and other various wild beasts of prey — towards the inside of the hedges until they were all in that paddock. The Emperor of Islam with Bulugan Khatun sat solemnly on the platform which had been built in the center and admired those animals. They killed some of them and set the others free” (p. 188–189, cited by Vereshchagin, 1959). It is worth saying that in the Middle Ages such entertainments involving a lot of human resources and cavalry were quite common among the aristocrats. It was the main hunting trick of many of the nomad tribes of that time living in Chapter 2. The Wisent in the Caucasus 19 the endless plains of the Western Asia and the South Caucasus. It is known that in 16th–17th centuries in Iran there was even a special hunting servitude for the peasants called “shikar” which they had to carry out from time to time. All this of course had impact on the preservation of the species in the Caucasian Isthmus, and as noted by N.V. Vereshchagin (1959), by the middle of the 19th century many species of large mammals had become extinct in the Caucasus: lion, tur, koulan, cheetah, beaver, elk, tarpan, tiger, etc. “Upon the order of shah Emir hunters with their henchmen and mulazims sometimes gathered up to ten thousand and more people for hunting, they en- circled a large area determined for the hunt with a special occlusion chain to prevent the animals from breaking the noose. This noose which was gradually tapering as the hunters were moving forward was called “jerke”, “negre” or “ko- marga” (p. 292, cited by Vereshchagin, 1959). During shah hunting in Minkale in the forests of Mazandaran (1618 A.D.) the noose was 3 farsangas (18–20 km) long and 2/3 farsanga (4–4.5 km) wide. Before the big hunting during which hundreds and thousands of large animals were killed the animals had been forced into the noose. Apart from the cavaliers they often involved infantrymen as drive beaters — “piyyadegan” from rapyyat. During big shah hunting in the forests and midwoods of Gilan (southern coast of the Caspian Sea, 1564 A.D.) about 10 thousand peasants were involved to drive the beasts into the “jerke”1” (Vereshchagin, 1959). There is information that wisents were found in the territory of Abkhazia before the 19th century (Radde, 1899), Arcangelo Lamberti in the 1700s wrote that he had seen a “wild buffalo” in Mingrelia (Turkin, Satunin, 1902). Snippets of information do not let us ascertain the nature of wisents’ distribution in the southern slopes of the Greater Caucasus Mountain Range; however, they prob- ably came here from time to time from the Kuban Region. It was also mentioned by N.Ya. Dinnik who wrote that in Abkhazia and on the coast of the Black Sea, in the headstreams of the biggest rivers — the Sochi River, the Shakhe River and the Mzymta River — he had not seen any traces of wisents. Describing that, the scientist also provided the results of survey of local hunters who had never met wisents in those forests but they claimed that their forefathers had hunted wisents in those places. In spite of the large habitat of the species in the past, by the beginning of the 20th century the wisents in Russia had been preserved only in the north-western part of the Caucasus, and they had lived there 100 years more than in all the remaining part of the habitat. There are different opinions about the sources and proofs of their life there. It is worth noting that in spite of the fact that the wisent is a large animal which was hunted in all the historical periods, the information 1 May be “jerge”, which translated from the Turkic means “into the ground” or “into the pit”, “pothole”, etc., author’s note. 20 U.A. Semenov The Wisents of Karachay-Cherkessia about it was provided only in the end of the 17th century. At that time Gulden- stadt who had visited the Caucasus in 1770 became the fi rst among the Rus- sian scientists who described wisents’ skulls which he had found in the places of sanctuaries in the caves of the Western Caucasus (Dinnik, 1910). However, the scientists obtained factual evidence for a more detailed description only 65 years later. In 1835 baron Rosen, Commander of the civilian service unit of the Caucasian corps of the Imperial army, sent a skin of a wisent to the Russian Academy of Sciences (Satunin, 1914). In 1838 A. Nordmann published the ar- ticle where he informed about the wisent’s spreading in the part of the Caucasus which was situated between the Kuban River and the Psib River source (Nord- mann, 1838). Academician Karl Baer compared the skin sent from the Caucasus with the skins of Lowland European bison (Bison bonasus bonasus) and made a conclusion that these species were quite similar and made the fi rst description of the Caucasian wisent. However, the provided proofs based on only one skin cast some doubt on the existence of wisents there until 1867, when the fi rst live wisent which had been caught in the Caucasus was brought to the Moscow zoo. Later on, in 1886 the Caucasian wisent calf caught by Adzhiev in the Verkhniy Urup (Karachay-Cherkessia) was brought to the same zoo. Comparative analysis of the documentary painting of this calf and the animals of the same age from the Bialowieza Forest carried out by M. Zablotskiy (1956) did not reveal any signifi cant difference. Similar conclusions were drawn in the earlier studies, as well (Wroblewski, 1927).

Figure 4. Caucasian Wisent. Painted by A. Koychuev. Chapter 2. The Wisent in the Caucasus 21 It is worth mentioning, that rumours saying that the wisents were living in the Caucasus were known long before the appearance of a live animal in the capital of the Russian Empire. Thus, in Decree No. 7994 of the cabinet of Her Majesty Empress Anna Ioannovna dated December 31, 1739 “On Annual Capturing and Sending of Various Live Animals to the Court and to the Izmaylovskiy Zoo” it was prescribed that the Chief Commandant of Astrakhan should send the ani- mals to the Izmaylovskiy Zoo, and in the amendment to this Decree it was writ- ten: “We also know that in Kabarda there are wild bulls and kdoses which are called “dombay” in their language1 and you have to exert all possible efforts sparing some expense from Our treasury in order to make local Dukes to order to capture and send to the Kizlyar Fortress young bulls and cows (fi ve or ten each), and there feed and lure them with bread and when they get used to this they shall send them to Astrakhan by water; and from Astrakhan they should be sent with other animals to Moscow, and in your own name you have to write to Shamkhal of the Mountainous Republic, to the Commandant of the Kizlyar Fortress and to Elmurza Cherkasskiy saying that they have to use the best endeavor to capture and send those bulls and cows” (Complete Collection of Laws of the Russian Empire, 1830). In addition to that, there is information that in December 1741 in the Izmaylovskiy Zoo there were already eight wisents, and it was planned to have nine animals in 1741 (“Internal Life of Russia”, 1880). P. Pallas in his “Zoographia Rosso-Asiatica” mentions the statement of acade- mician Lovic saying that he captured a large wisent in 1776 in the headstreams of Kuma River not far from Beshtau Mountain, and that academician Anton Gul- denstadt found a skull of that animal in the caves along the Urukh River Valley (Pallas, 1831; cited by Kirikov, 1979).

2.3. Morphology and External Appearance of the Caucasian Wisent

Today it is not possible to provide precise morphological characters of all the sizes of the Caucasian wisent. In the foreseeable future rampant development of digital technology will most likely let us recreate precise models of animals on the basis of separate fragments of the bodies available in the scientifi c archives of the country. In spite of the fact that the Caucasian wisents had been exter- minated before the scientists had time to study them more or less in detail, in literature there were bits of information about some measurements made by the

1 “Dombay” is translated as “wisent” from the Karachay and Balkar dialects which are in their essence the language of one and the same people divided by Elbrus mountain into the Kara- chays and the Balkars, but at that time the lands of Karachay and Balkaria were little known to the general public because they were hard-to-access, and the peoples were small-numbered, that is why these lands were more often referred to as part of the other lands and under the name of other bigger ethnic groups under the unifi ed name of “Kabarda”; author’s note. 22 U.A. Semenov The Wisents of Karachay-Cherkessia scientists in the end of the 19th and the beginning of the 20th centuries (Shilder, 1901; Dinnik, 1910). Many researchers expressed the opinion that the Caucasian wisents having certain adaptations to living in the mountains are smaller than their congeners living in the plains (Heptner et al., 1961; Flerov, 1979), however, as remarked by A. Danilkin (2005), those conclusions turned out to be quite doubtful when the source materials which had been used by the natural scientists were studied more in detail. There are too little samples of the Caucasian wisent, the studied param- eters are very variable, and their application has low accuracy due to the usage of data about animals of different age. Earlier other researchers also expressed doubts related to that matter (Bohlken, 1967; Sokolov, 1971), and asserted that there was not enough evidence to consider the Caucasian wisent to be a smaller animal. Already in the beginning of the 19th century N.Ya. Dinnik (1910) noted that there was very little reliable information about the size of the Caucasian wisent. According to his data, one of the fi rst measurements were made by G. Shilder who in 1895 hunted in the mountains of the Kuban Hunt with Grand Duke Ser- gei Mikhailovich. Then they got an old female wisent which was 4 arshins 5 vershoks (3.7 m) long from nose to tail. Distance between the bases of horns was 4 ½ vershoks (20 cm), distance between the horn tips was 1 ½ vershoks (7

Figure 5. Native Caucasian Wisent (Old Bull) Captured by D. Filatov in 1909 in the Head- streams of the Kishi River (Filatov, 1910). Chapter 2. The Wisent in the Caucasus 23 cm), circumference of the horn base was 4 vershoks (18 cm), circumference of the chest was 2 arshins 14 vershoks (2.7 m) and length of the foreleg was 18 ½ vershoks (82 cm). In the same year they obtained a bull whose head-and-body length was almost 3 m, distance between its bases of horns was 24 cm, distance between the horn tips was 28 cm, circumference of the horn base was 28 cm, horn length was 37 cm, height at the shoulder was 1.6 m, length of the lower margin of hoof was 14 ½ cm. Further in his study he provides information about mounts of the male and fe- male Caucasian wisents whose proportions were as follows: height at the shoul- der — 1.56 m, at the sacrum — 1.4 m, head-and-body length — 3.79 m, head length — 56 cm and height at the shoulder — 1.42 m, at the sacrum — 1.3 m, length — 2.6 m, respectively. The scientist himself pointed out that the horns of the Caucasian bulls can be signifi cantly thicker and larger. For example, cir- cumference of the horn base of the old bull was 33 cm, circumference of the horn base of the others — 35 and 31 cm, and their length varied from 33 to 42 cm (Dinnik, 1910). Nineteen wisent skulls which were discovered in the Olisay- dom Cave and thoroughly measured (almost all of them had forehead width at the rear end of the eye-pit varying from 29 to 32 cm), let him conclude that their size was almost the same as the size of skulls of the wisents inhabiting moun- tains of the Kuban Region. As noted by the author, severe damage of the facial part of the skulls did not let him collect detailed data about the length of the majority of the discovered bones, but in two cases he managed to do it more or less accurately. The length of the fi rst skull was 56 cm, of the second one — 54 cm. N. Dinnik (1910) also stated there that the skulls found by Guldenstadt were signifi cantly smaller in size, and their length was 40 cm, and the width between eye-pits was 21 cm. General body build of the wisent is large and heavy, typical for bulls. Male spe- cies are bigger than female species and differ from them in a larger front part of the body. The body length of modern male animals at the average is 281–289 cm (maximum up to 332 cm), height at the shoulder — 180–183 cm (maximum up to 210 cm), and body weight — 615–631 kg (maximum up to 920 kg). Female species’ size and weight are smaller than those of male species: body length — 240–262 cm (up to 290 cm), height at the shoulder — 169–174 cm (up to 197 cm), body weight — 424–439 kg (633 kg). The tail is long, up to 92 cm, with hair at the end. Ears are relatively short and wide, normally they are not longer than 16–18 cm. Eyes are not big and partly protruding. All the wisent body is covered with fur which is especially thick and long on the chest and neck. Above the neck and on the withers there is a mane of slightly wavy over hair. Summer coat is thinner and shorter (Sokolov, 1959; Kalugin, 1966; Kiseleva, 1974; Nemtsev, 1988а; Sokolov, Tembotov, 1993; Sipko et al., 1999, 2002; Sipko, 2002). When providing data on the wisents’ size some scientists state the record values for certain animals. For the 24 U.A. Semenov The Wisents of Karachay-Cherkessia European bison (Bison bonasus bonasus) it is 1,900 kg, for the native Caucasian wisents it is over 1,000 kg (Bashkirov, 1939, 1940). May be such animals could have been found in the past when the wisents’ habitat was large, and their historical degradation in size happened due to the worsening of living environment condi- tions (Kulagin, 1919). The native Caucasian wisent was less heavy and had more developed pectoral girdle. It is considered that this is animals’ adaptation for living in the mountains where forelimbs of the animal bear additional load while going down the slopes (Nemtsev et al., 2003), though the bison living in the open spaces have similar body structure. Unfortunately, we have little information based on which it is pos- sible to estimate the body size of the Caucasian wisents, sample amount for the Caucasus before 1921 is only one to three species. Nevertheless, they provide a certain understanding of real size of the animals which lived there before their extermination: body length with the tail — 307 cm, height at the shoulder — 160 cm, circumference of the chest — 207 cm, body weight — up to 1,000 kg, horn length — 33–42 cm (Dinnik, 1910; Danilkin, 2005).

2.4. The Wisent in Karachay-Cherkessia

One of the fi rst references about the wisents in Karachay-Cherkessia was made in 1864 by baron F.F. Tornau, a Russian intelligence offi cer, who undertook a diffi cult and dangerous trip from Abkhazia to the Kuban Region together with his Abkhazian guides. In his memoirs called “Memoirs of the Caucasian Offi - cer” he mentioned several times that the wisents existed in those places: “When Mikambai remarked that we did not have enough food for such a trip, Khatkhua objected to him saying that with the shot-gun it was impossible to die of hunger in the places where there were goats, turs and adombeys” (Tornau, 1864, p. 89), further on F.F. Tornau (in the same story) explained that “adombey” was the Ab- khazian name of the wisents which lived in the northern slope of the Caucasus in deep chines near the headstreams of the Urup, the Bolshoy Zelenchug and the Maliy Zelenchug1. Describing the place of his trip he wrote: “To our right and left there were crys- tal white mountain battlements. There were black Abkhazian mountains behind, and the sea was seen in the distance as a blue line. In front of us we could see cliffs, forest-clad edges and green grass-covered spurs of the Greater Mountain Range, and between them the Kuban River and rivers fl owing into it gleamed like silver ribbons. The weather was wonderful, there were no clouds in the sky, there was no wind in the air, it was silent all around, everything near us and in the distance was lit with glaring light. The sight of the Black Sea and at the same

1 Today it is the Zelenchuk River, author’s note. Chapter 2. The Wisent in the Caucasus 25 time of the Caucasian steppes from the height of eleven thousand feet above the sea level was startling in its splendor. The Greater Mountain Range col which we crossed lay between the headstreams of the Bzyb running into the Black Sea and the Bolshoy Enjik-su running into the Kuban. The Russians converted the word “Enjik-su” into a resembling one “Zelenchug”” (Tornau, 1864, p. 94). Trying to restore the itinerary of the intelligence offi cer we can say with great probability that Tornau with the guides started from the headstreams of the Bzyb which was located in the north-west of Abkhazia, crossed the mountain pass1 of the Greater Caucasus Mountain Range, and came into Karachay-Cherkessia, to the headstreams of the Amanauz and the Psysh. Confl owing with numerous feeders they form the Bolshoy Zelenchuk River downstream. The detailed de- scription of his itinerary seems to be important because there the group met a Caucasian wisent for the fi rst time. “…In a few minutes we heard a fast-ap- proaching brattle mixed with the sound of crashing trees broken by some insu- perable power in the forest. Then, the disturbed herd of adombeys came in front of us within a good shooting distance, a huge bull was running ahead of them. The guns were immediately taken out of the covers, the shots thundered, and the bull at which we all targeted made a high jump and turned to the opposite side. The whole herd which contained up to twenty cows and calves followed the bull” (Tornau, 1864, p. 97). During that incidental hunt, which happened to the travelers, they heavily wounded an adult bull with four bullets which had passed clear, and on the next day they killed it with seven more shots at the watering hole. Tornau remarked (1864) that in spite of the signifi cant blood loss the wisent had stayed alive till the morning and even moved. Understanding the importance of that incident the baron made an attempt to preserve the wisent’s skin and present it to the Russian scientists, but the necessity to move fast and covertly made him refuse from that idea. He wrote ruefully: “I wanted to preserve the adombey’s skin with its head and hoofs and bring it to the Line as a proof that the wisents lived not only in Lithuania but also in the Caucasus, which was not believed in, though we heard about it from the mountain people. I was the fi rst among the Russian people who had the possibility to see a Caucasian wisent and hunt it. Its huge height, dark- brown color, purely bovine head as hairy as its chest and smooth hinder part did not let me doubt that it was of the same breed as the animals preserved in the Bialowieza Forest. In the Caucasus the wisent is found in the pine woods verging the permanent snow, in the chines of two Zelenchug Rivers, the Kyafar and the Urup. Several times during the journey they showed to me lairs of those animals and the paths which they had made in the most arduous mountains and which were used further on to the benefi t of humans because they always led to some

1 Probably the Naur or the Medvezhiy, author’s note. 26 U.A. Semenov The Wisents of Karachay-Cherkessia chine creek where adombeys drank water. My Abkhazian people did not want to hear anything of taking the whole skin to the Line saying none of our horses had power to carry it for even one day and we would have to leave the horse on the road together with the skin. Against my will I had to obey to this rather prag- matic conclusion of my guides and I saw with disappointment how they made straps out of the perfect skin under which all the seven of us had found shelter during our stop on the bank of the Zelenchug” (Tornau, 1864, p. 100). This story of F.F. Tornau where he mentioned the herd of twenty animals and the presence of the youth of different age says enough about the satisfactory con- dition of the Caucasian wisent population in Karachay-Cherkessia in the middle of the 19th century. Taking into account that afterwards in the said places the wisent disappeared much earlier than in the headstreams of the Belaya River, the Bolshaya Laba and the Malaya Laba Rivers, we can suppose that at that time there were still many wisents in the Caucasus. It is impossible to determine pre- cise borders of their spreading within Karachay-Cherkessia because there are no data about the number of the Karachay chines in the headstreams of the follow- ing rivers: the Makhar-su, the Uchkulan, the Dzhalpak-Kol, the Ullu-Kam, the Khudes and others. Bits of information that we have managed to collect show that before the middle of the 19th century the wisents were found almost in the whole mountain part of Karachay-Cherkessia. One of the fi rst scientifi c articles about the Caucasian wisents in Karachay- Cherkessia was written by V. Latsarius in 1879, at that time the scientist de- scribing their habitat indicated that the wild wisents were found on the northern slopes of the western part of the Caucasian Mountain Range, in the southern strip of the headstreams of the Kyafar, the Aksaut, the Zelenchuk, the Marukha and the Urup Rivers through to the Malaya Laba River. At that time there were no big settlements, and a huge area was covered by untrodden broad-leaved and coniferous forests. Only in summertime some parts of the chines listed above were occupied by the shepherds with their numerous fl ocks of sheep and herds of horses from the mountain villages of the Batalpashinsk District of the Kuban Region which at that time comprised all the Karachay settlements. At the other time the chines were almost free from human activity, only the most reckless mountain people came that far driven by the hunting instinct. Diffi cult access to the forests and absence of roads became the main reason why the wisents survived there for a bit longer period of time while in the major part of the Cau- casus they had already been exterminated. Nevertheless, general decrease of the species habitat continued, and as stated by V. Latsarius (1879), it was possible to see a live wisent only if you took great care, hid or if you lay in ambush before- hand waiting for the wisents near watering holes or passage points. Probably the scientist had never met wisents himself because he described their exterior based on the Cossacks’ stories. As mentioned in the article, the Caucasian wisent bore Chapter 2. The Wisent in the Caucasus 27 resemblance to the European bison (B. b. bonasus), its coat was rusty-brownish without any speckles. Male species was markedly larger than the female one and could be quite furious when enraged. The herds amounted up to 30 species and had their leaders. Then he described the animals’ behaviour: “The wisent has quite poor eyesight. If you fi nd wisents and stay in such an ambush from which they cannot smell the humans, you can shoot at them several times, and they will not run away or towards the shots, they will only smell the fallen victims in surprise; and only after repeated shots the herd will slowly go away, and it seems they do it not because of danger but because they are afraid of powder fume smell. If the wounded animal sees its enemy it becomes enraged and, of course, the blow of its mighty horns is mortal” (Latsarius, 1879, p. 7). Describing the snowy winters the scientist provides rather interesting facts saying that the wisents looking for food went down into the valleys where they fed on tree branches and lichen, and the starving and weary animals came close to the settlements looking for hay laid out on the branches where they were ruth- lessly shot with impunity sometimes from a very short distance. And all that hap- pened in spite of the imperial prohibition against shooting wisents. In particular, many such cases were noticed near the Storozhevaya Cossack Village. N.Ya. Dinnik (1884, 1890а, b, 1899, 1909 and others) who had studied the headstreams of the Karachay-Cherkessian rivers for a long time became one of the fi rst researchers of fl ora and fauna of those mountains, his vivid and interest- ing descriptions left a deep imprint on the studies of the Caucasian nature, and the fi rst research parties of 1875–1877 went to the highlands of the north-western part of the region. Descriptions of animals and birds made by Nickolay Yakov- levich Dinnnik during his trips along the valleys of the Teberda, the Dombay- Yolgen, the Bu-Yolgen, the Zelenchuk, the Zagedan, the Bolshaya Laba, the Ma- laya Laba, the Urushten, the Belaya, and other rivers, give the possibility to look back and see what the Karachay-Cherkessian fl ora and fauna looked like two centuries ago. As N.Ya. Dinnik devoted all his lifetime to studying the nature, he went through the wildest places of the mountains and chines of Karachay1, and that let his contemporaries perform comparative analyses not only in the sphere of animal wildlife but also in glaciology, botany and fl oristics. In 1890 in one of his fi rst articles in The Priroda i Okhota Journal he wrote: “…headstreams of the Laba and the Urup are without doubt the richest places in game animals in the whole Caucasus” (Dinnik, 1890, p. 45). During one of those journeys Dinnik together with local hunters explored the Teberda Chine and passed through the valleys of Bu-Yolgen and Dombay- Yolgen. “Above the place where the Khadzhibey disembogues into the Teberda River the road, however, became signifi cantly worse. Near the river bank a nar-

1 At that time Karachay was a part of the Kuban Region, author’s note. 28 U.A. Semenov The Wisents of Karachay-Cherkessia row path completely overgrown with hazel and other shrubs ran towards the side-hill. We had to walk a certain distance up the path. About two o’clock in the afternoon we reached the place where the Teberda River turned to the south-west at a right angle and then continued its way in its arm, the Dombay-Olgen” (Din- nik, 1884, p. 8). There he explained: “Dombay” — “wisent”, “olgen” — “killed” or “wounded”. “Probably wisents lived there in the past”, the author wrote1. The same names of the mountain, chine and river, Dombay-Yolgen, were translated by the scientists in different ways, but the meaning was more or less the same: hunted or killed wisent. Though there are reasons to believe that this is not exact. Taking into account that Dombay-Yolgen is quite an ancient name (in 2011 Khasan Bostanov, born in 1907, one of the oldest residents of Kara- chay-Cherkessia narrated that his father and he together with other people of Kyzyl-Kala mountain village had driven cattle to the summer grazing in Dom- bay and Teberda. At that time the main catstep ran along the left bank of the Teberda River which branched off to the subalpine and alpine zones towards the alpine meadows where the summer grazing areas (koshi) of the mountain people were located. He remembered that the elders said that even their forefathers had known about those places and had come there in small groups, presumably it was the end of the 16th century and the beginning of the 17th century. At that time wisents were not rare, and capturing of such a common animal could hardly be- come such a remarkable event to name mountains, chines or rivers after it. Most often such names were not created intentionally but as a result of certain inci- dental or remarkable events. Moreover, the word “yolgen” (“olgen” or “ulgen”) literally means “dead” and not “killed”. Probably a wisent which had no time to leave the valley because of the fallen thick snow got into an avalanche or died here because of a wound. After that it was discovered by shepherds or hunters and served as a sort of a guide mark for them when indicating the place. The name was vernacularized and then spread. With time a mountain mass resem- bling a powerful wisent body (town of Dombay-Yolgen — 4,046 m above the sea level), a mountain stream (the Dombay-Yolgen River), and an isolated ter- rain got the same names. Otherwise, the name would contain the word “yoltur- gen” (“Dombay-Yolturgen”) which would mean “killed” (“hunted”) wisent, and not “yolgen”. Anyhow, the name expressly confi rmed that the wisents had lived there before. However, in 1884 when N.Ya. Dinnik explored those places he did not fi nd either those “forest giants” or their traces there. In the same year he explored the chines neighbouring with the Teberda Chine along the valleys of the Marukha and the Aksaut where he did not fi nd either wisents or their trace fossils.

1 Different scientists quite often wrote down the local names by ear and that of course led to existence of different variants mentioned in the literature. In relation to that place there were also “dommay”, “ulgen”, “yolgen”, author’s note. Chapter 2. The Wisent in the Caucasus 29 The next expedition of the natural scientist in 1884 was directed to the head- streams of the Urup and the Laba: “In the headstreams of the Laba River there is the Zagedan Valley, wonderful in many ways. It is possible to get to it only by barely noticeable and often very bad mountain paths; therefore, people seldom come there, and it is basically wild and undisturbed nature, where there are still thick forests inhabited by various wild animals, and no man has ever set foot on many places of those forests, and the sound of axe has never been heard there. As we will see below, nowadays, these forests are inhabited by beavers (Cas- tor fi ber), wisents, which are exterminated almost everywhere, and many other animals. I wished to visit the Zagedan as the most interesting place from the zoo- logical point of view. I hoped to meet turs, chamois, deer, mountain blackcocks, and perhaps wisents or beavers” (Dinnick, 1884, p. 21). When Dinnik was in the headstreams of the Urup and the Bolshaya Laba Riv- ers he did not manage to see the wisents but he discovered three strides of those animals. Polling data which he carefully collected inquiring all the met hunters, shepherds, and analyzing the data in the articles about the Caucasian animals, let him get an overall picture of the geographical range of the Caucasian wisent in the Karachay and the neighbourhood of the Kuban Region as of the end of the 19th century. “In the recent years there have been no wisents during summer in the head- streams of the Urup, so at this time of the year it is impossible to meet wisents to the west of the valley of the Laba and its tributary called the Pkhi. At the same time according to the information from the Kuva hunters, the wisents can be found in the headstreams of the Urup in winter. Byty, our guide, who without doubt is a good hunter, killed fi ve wisents in that area several years ago. The Karachays claim that the wisents come to the headstreams of the Bolshoy Zelen- chuk, in particular, to the Irkyz Valley, and as the latter is separated from the forests with a strap along the Pkhi River which is only two or three versts long, sometimes the wisents certainly get to the Irkyz [Dinnik explains that: “The Irkyz is a thick forest in the headstreams of the Bolshoy Zelenchuk. It is very similar to the Zagedan” (Irkyz or Yrkhy-yz1); but the forest cannot be considered a permanent habitat of the wisents” (Dinnik, 1884, p. 52)]. The way of life of these animals has always been characterized by seasonal and daily migrations favoured by infl uence of various factors. For instance, as indicated by Dinnik, when spring and summer came wisents left the Zagedan trying to hide from heat and gnats, and migrated to remote and wild areas in the headstreams of the Bolshaya Laba. At that time there were no shepherds and settlements there, only hunters could reach those places looking for exciting hunt and enviable spoils of chase. Coming to those places they witnessed a rich

1 Trace of mudslide or mudfl ow, was at that time the name for Arkhyz, author’s note. 30 U.A. Semenov The Wisents of Karachay-Cherkessia variety of fl ora and fauna where the wisents, “forest giants”, could be easily met when they came out to the forest glades and made wide paths in the forestlands. But when autumn came, and the temperature went down, the wisents gradually moved downstream back to the Zagedan and along the Urup. During especially snowy winters the animals had to get even lower appearing in the forests sur- rounding the Cossack villages. In such winters the wisents were even hunted not far from Storozhevaya Cossack village (Dinnik, 1884). Describing the wisent range in the upper part of the Kuban Region Nickolay Yakovlevich Dinnik wrote: “In the past the wisents probably lived farther to the east, particularly in the valley of Teberda. As the Karachay people say this is proved by the name of the Teberda tributary, the Dombay-Olgen, meaning “wi- sent kill” in Russian (Dinnik, 1884, p. 52), but he disagreed with the opinion of V. Latsarius that wisents also lived in the headstreams of the Marukha and the Aksauta, and he stated that they had not lived there for 40–50 years already. He also added that the conditions for their life there were quite doubtful because there were no wide forests where they could live. Comments of N.Ya. Dinnik to the article of V. Latsarius (“Article about the Caucasian Wisent (Bos urus L.)”) disprove the opinion on wisents’ spreading in the chines mentioned above. However, it is still unclear when V. Latsarius visited Karachay, and to which years the information published in 1879 is re- lated. Nevertheless, about fi ve years after the publication of V. Latsarius wi- sents’ habitat in Karachay-Cherkessia underwent signifi cant changes. Settlement in the chines and their intensive exploration by humans were rapidly reducing the original habitat of those animals. We can suppose that it was not possible to meet the wisent herds consisting of up to 30 specimens in those places, as stated by V. Latsarius, but small groups or solitary animals could have come to those places till the beginning of the 20th century. Studying the western borders of the wisent range Dinnik wrote that those bor- ders went on through the headstreams of the Malaya Laba, the Khodz to the Belaya River, probably to the source of the Pshekha River. “I was told that in the huge forests surrounding Abago mountain situated not far from the source of the Belaya, there were as many wisents as in no other place in the Caucasus. If we consider that the wisents’ range lies from the Irkyz to the sources of the Belaya River, its length will amount to 85 or 90 versts, and its width, even if we take into account their migrations, will amount to 35 or 40 versts in the eastern part and even less in the western one. I do not know anything exact of wisent existence on the southern slope of the Caucasian Mountain Range. Some people (for example, Nordmann) claimed that it lived there (as for example in Abkhazia), but the Karachay hunters stated that the wisents lived only on the northern slope” (Dinnik, 1884, p. 53). Five years after Dinnik came to Karachay-Cherkessia again, that time he ex- plored the headstreams of the Bolshoy Zelenchuk and the Abishara-Akhuba Chapter 2. The Wisent in the Caucasus 31 Mountain Range, his main goal was to discover wisents in those places. His search was not successful, at that time wisents were not found there already. In the conclusion of his article he wrote: “My personal observations and inquiries of many hunters and forest rangers persuaded me that in spite of assurances of some modern authors at that time the wisents were found neither in the head- streams of the Bolshoy Zelenchuk nor in the headstreams of the Urup. I can also say the same about the valleys of the Marukha and the Aksaut which I visited earlier. The headstreams of the Zelenchuk and especially of the Irkyz (Arkhyz) are very comfortable for wisents’ living but they left those places about 35–40 years ago. Probably they were ousted from that area by the forest felling and mountain shepherds with their baranta and herds. In the headstreams of the Urup covered by vast continuous forests the wisents stayed for a relatively long time, in particular, till the beginning or even the middle of the 1880s. Farther to the west we see the place where the wisents live nowadays. When travelling to the headstreams of the Laba in the mid-80s I saw a lot of wisent paths, their fresh traces and even live wisents but now wisents come to these places only from time to time. More often they can be met along the valleys of some tributaries of the Malaya Laba, for instance the Alous, the Mastakan (Mestyk), and especially the Urushten, but even more wisents live in the valleys and chines of numerous sources of the Belaya River, in the terrain which is completely free of humans and covered with thick forests. Farther to the west in the headstreams of the Pshekha the wisents do not live already. Thus, we see that for the last 30–40 yeas their range has narrowed signifi cantly particularly from the eastern side” (Din- nik, 1899, p. 38–39). The next expedition of the natural scientist in 1903 was dedicated to explor- ing the headstreams of the Bolshaya Laba and the Tsagerkel Mountain Pass. The main goal was to explore the isolated terrains of Damkhurts, Mamkhurts and Makera along the beds of the rivers of the same name to see whether there were wisents. Before 1885 those places constituted thick impenetrable forests where nobody except for the shepherds and hunters could get to. By the time when the expedition was taken the road had been laid there, veterinary stations had been established, and forest felling had been started. Dinnik wrote: “In the past there were many deer, bears there, not to speak of lynxes, martens, otters and big ani- mals; even wisents lived there, which I personally checked on during my far-off trip to that part of the Laba which was known under the name of Zagedan, and whole herds of turs and chamois walked in the mountains surrounding the chine of Laba. Now the majority of wild animals have migrated from these places. Only valleys and chines of the abovementioned tributaries of the Bolshaya Laba have stayed in the primeval condition, and nowadays there are neither roads nor even paths, and that is why it is not possible to get there with horses and the nec- essary store of food” (Dinnik, 1905, p. 2). 32 U.A. Semenov The Wisents of Karachay-Cherkessia Exploration of those isolated terrains which in the past had belonged to the most remote and deserted places of the time in the whole Caucasus, showed total absence of wisents and their traces there. A similar situation was in the neighbouring chine in the valley of the Sanchara River. At the same time there were many animals there, the abundance was proved by Dinnik’s article (1905) where he wrote that two huntsmen of the Kuban Hunt of Grand Duke Sergei Mikhailovich had killed more than ten bears for four days there. Analyzing the literature about the wisent occurrence in the Caucasus starting from the mid-18th century till the beginning of the 20th century we can conclude that they inhabited all the northern slopes of the Greater Caucasus Mountain Range. However, the natural habitat of the species was rapidly decreasing, and in the beginning of the 20th century the wisents stayed only in the small area start- ing in the headstreams of the left tributaries of the Kuban River, in the basin of the Malaya Laba River and ending in the basin of the Belaya River in that part which was rented by His Imperial Highness Grand Duke Sergei Mikhailovich for hunting. Ideas expressed by some authors with regard to the wisents’ occur- rence on the southern slopes of the Greater Caucasus Mountain Range, and in Abkhazia, were fallacious according to N.Ya. Dinnik’s opinion: “I think that the opinion expressed by Radde and supported by some other authors with regard to the fact that wisents are found in Abkhazia and go down almost to the Black Sea is completely fallacious. Neither in the headstreams of the Sochi, nor the Shakhe, nor the Mzymta, nor the Kodor I have ever seen their traces, and all the hunters and forest rangers of those places told me in the same breath that wisents did not live there at all. But once they told me that wisents’ traces had been seen upstream Romanovsk (today Krasnaya Polyana urban locality), but as seen by the traces the wisents stayed there for a short period of time and, probably, re- turned to the Kuban Region. About 25 years ago the Karachay hunters who often had to get to the southern slope of the Caucasian Mountain Range in the district of Sukhumi and in the former Black Sea district told me that wisents did not live there. As has already been said, the Abkhazians told the same thing to baron Tornau” (Dinnik, 1910, p. 147–148). In the end of the 19th century the Caucasian wisents had to be sent to the Bi- alowieza Forest and because of that hunters were sent to the indicated region to capture young animals. This activity involving additional forces turned out to be quite a large-scale operation and helped them to determine approximate quan- tity of the native wisents in that region. While looking for newborn calves the hunters noticed several herds consisting of 12–50 animals. It is remarkable that the hunters found a herd of 20 species in which almost all the animals were old bulls in the region of the Alous and the Urushten. Moreover, one of the huntsmen of the Kuban Hunt informed Dinnik that he had seen a herd of 27 wisents near Slesarnya. Chapter 2. The Wisent in the Caucasus 33 “Taking into account, on the one hand, the diffi culty to access this terrain, thick forests, tall wild grasses growing there in summertime when hunters looked for wisents, due to which only a small part of them was seen by the hunters, and on the other hand, that fact that the wisents were not looked for in the Kishi Valley where the frequency of their occurrence is higher than in all other places, we can think that there are at least several hundred wisents, most likely 500–600 animals in these places. The wisents do not like solitary life. Even old bulls rarely live solitarily, usu- ally they live in groups of two or four animals. Most often wisents stay in small groups of six to eight species. Such a group usually consists of young cows (aged two to three years), calves and old cows. Sometimes the wisents gather into big- ger herds. About seven years ago huntsman Chugunov saw a herd of 18 wisents near Slesarnya; in that herd there were two old bulls, about ten cows and almost the same number of calves aged from one to two years. In 1900 people saw a herd of 14 animals consisting of old cows, calves and more or less adult bulls. Not old bulls also often live in groups of six to eight species. Nobody has ever seen very large wisent herds in the Caucasus” (Dinnik, 1910, p. 149). Thus, in spite of constantly decreasing habitat of the Caucasian wisent, it was still preserved in the beginning of the 20th century. Complicated political situ- ation of that time partially helped to decrease the hunting pressure on wisents. Mountain peoples pressed back to the remote diffi cult-to-access areas during the Caucasian war almost did not arrange any settlements in the places where wild animals dwelled in wintertime, in the mid-mountain part of the spurs of the Greater Caucasus Mountain Range. Those wide and little-inhabited places were favourable for animals’ foraging and wintering, and in summertime when there was a lot of food animals found the possibility to move up the mountains by numerous animal paths. By the mid-18th century the Russian Empire had made a lodgment in the steppes of the North Caucasus, forming the Caucasian Line from the source of the Terek to the source of the Kuban, and signifi cantly expanding its southern borders. Creation of the boundaries and conquest of the Black Sea line reduced the stress of warfare, some peoples were resettled to the Urup, part of them mi- grated to Turkey. Such politico-military operations changed a bit the location of mountain people’s settlements which returned to the mid-mountain and high- mountain parts pushing the wisents back to the upper parts of the chines again. Starting from 1851 active situation shifted to Kuban, and in 1864 all the Western Caucasus was under control of Russia. Cessation of hostilities let the mountain people return to their normal life, and the hunting pressure on wild animals, in- cluding wisents, started to grow intensively. Such a large animal as wisent found it more and more diffi cult to fi nd quiet corners for living, and the only thing that saved wisents from complete extermination for some more time was the organi- 34 U.A. Semenov The Wisents of Karachay-Cherkessia zation of the fi rst protected natural area in the Caucasus called the Kuban Hunt of the Grand Duke. It was created upon the initiative of the Grand Dukes of Ro- manov dynasty in the territory of 522 thousand dessiatines in the forest dachas of the Ministry for State Property and in the lands transferred to the Cossacks of the Kuban Regional Military Board which had belonged to the mountain people before. Borders of the Kuban Hunt of the Grand Duke lay along the Greater Caucasus Mountain Range, where the border between the Kuban Region and the Black Sea district was located, in the south; along the Bolshaya Laba within the present-day borders of Karachay-Cherkessia in the east; along the Belaya River in the west; along the Peredovoy Mountain Range in the north. Huntsmen were specially hired, money was allocated for protection of the territory, control over vermin, improvement of facilities, etc. in order to arrange imperial hunts there. It is worth noting that the wisent hunting was prohibited in the whole territory of the Russian Empire, and Grand Duke Sergei Mikhailovich hunting there several times a year had to obtain a permit for the wisent hunting from “His Highness”. The place for organization of the Grand Duke’s hunting was chosen know- ingly by F.I. Kratkiy who knew those places very well and was in charge of the Caucasian hunts, and in 1898 N.Ya. Dinnik travelling along the Urushten and the Belaya got to the headstreams of the Kishi to defi ne the wisent’s condition, and thoroughly explored that area, the situation with preservation and the variety of fauna. Admiring one of the most extensive and rich sectors called “dark spruce forest” he wrote: “A long time ago Sakhray hunters (nimvrods) hunted deer, wild boars, bears there and killed wisents many times, but now when the watch sta- tion for 3 huntsmen of the Grand Duke’s Hunt is built on the Kishi, it has become impossible for them. …It is the whole world of the deepest chines, valleys and draws, covered with thick forests where humans have hardly ever set their foot. But we can say they are full with various game animals: deer, bears, wisents, wild boars, lynxes, etc. Hunters of the Grand Duke who reached those places last year saw not less than one hundred deer, and killed a panther and four bears in one day” (Dinnik, 1898, p. 36). “I was told that this forest was one of the favorite wisents’ places, and that is why I especially wanted to see it. To get to the dark spruce forest we had to descend from Slesarnya mountain and cross the narrow glade overgrown with wild grasses in some places, and with marsh plants in the others. At the edge of the glade we saw a spring very often visited by the deer and sometimes by the wisents. Some time ago the Sakhray people arranged there a deer blind in the form of a small booth hidden between the trees to be on the watch for animals. Not far from that place at the very edge of the wood we came across a huge fi r under which there was a wisent lair. As it is known wisents like to rest under secular fi rs with branches and fi r needles so thick that even the heaviest rain cannot brake through them. Soil in such places is always dry and covered with Chapter 2. The Wisent in the Caucasus 35 a layer of fallen fi r needles, and grasses never grow there because of the high level of shadowing. Under another fi r there were two large wisent lairs. Farther in the forest we saw a huge fi r torn up by the root near which wisents had lain, dug the ground with their hoofs and rested many times. In the territory which was 4 sazhens long and wide the ground was dug up, loosened, trampled down again and was completely free of grass. Farther under the group of gigantic fi rs we saw six wisent lairs. It could be noted all around that wisents were constantly dwelling there, and that there were quite many of them. This was the situation in 1892. Now due to increased protection of this area from poaching there may be even more wisents here” (ibid., p. 37). Organization of the Kuban Hunt of the Grand Duke certainly played a key role in the preservation of the Caucasian wisent. Isolated and well-protected extensive territory practically guaranteed successful preservation of the animals there for a long time. However, the events of 1917 in Russia, and the chaos and civil war following them not only deprived the Caucasian subspecies from the opportunity to survive but also laid down wisents’ existence at stake. Before the beginning of the 1st World War dated 1914–1918, about 785 wi- sents lived in the Bialowieza Forest (Severtsov, 1926, 1941; Zablotskiy, 1947), descendants of wisents from the Bialowieza Forest were kept in the Pshinskiy Hunting Park of Prince of Pless (Poland), there were 74 species in total brought there in 1865 and 1893. Additionally, wisent male species were brought there from the Berlin and Dresden Zoos in 1883. In 1917 in the Gatchina Imperial Hunt (near St. Petersburg) there were 36 wisents, 22 animals were in the Hunting Park “Pilyavino” (Ukraine), and nine wisents were in the lands of the Crimean Imperial Hunt. A small number of animals also lived in zoos and parks of other countries. Thus, at that time there were about 1,000 European bison (Bashkirov, 1940), and not more than 500–700 Caucasian wisents (Dinnik, 1910, Filatov, 1910, 1912). Therefore, the world livestock population of wisents amounted to 1,500– 1,700 animals in the beginning of the 20th century. The 1st World War and the revolution following it in certain countries, which in Russia escalated into the civil war, led to disastrous decrease of the species population. So wisents had been exterminated in Pilyavino by 1917, the animals had disappeared in the Gatchina Hunt by 1918, in the Crimean lands they had been killed by 1919, and the last wisent in the Bialowieza Forest was killed on February 9, 1921. Only three wisents survived, including the animals called “Planta” and “Plebeier” (afterwards they were found in the wisent studbooks of the USSR), from the famous herd of the Prince of Pless. Diffi culty to access mountain areas, delayed aftershocks of new ideas of the revolution and the civil wars hampered a little the impact of the hunting pressure on the Caucasian wisents. In 1924 the newly created young Soviet country even 36 U.A. Semenov The Wisents of Karachay-Cherkessia organized a special wisent nature reserve to conserve the species. However, they did not manage to save the Caucasian wisents. After October events a lot of people with cattle, wood choppers and hunters came to the territory of the former Kuban Hunt, and in 1919 epizootic disease most likely brought in to the mountain grazing by domestic cattle burst out there. Poaching which had enhanced dramatically after the revolutionary events in the country continued even after the nature reserve had been organized. In the be- ginning of the 1920s the Caucasian wisents were not met anymore in Karachay- Cherkessia. It is not known whether the animals came to the headstreams of the Bolshaya Laba left bank. According to some statements (Bashkirov, 1939), the last three Caucasian wisents were killed on Alous mountain in 1927. Thus, by the time when the administration of the nature reserve had got the security service on the right track there were no more wisents left there, and it became impossible to restitute the thoroughbred race of the Caucasian wisents. The wisents have always attracted attention of hunters and natural scientists. They were captured, transported, and kept in different countries by separate per- sons of high standing and by special institutions (imperial zoos, hunting areas, parks, etc.). The Caucasian wisents were also exported from the Caucasus many times, and some of those movements of the animals were kept in the documents, and it became possible to retrace their destiny. In 1866 a small calf captured in the Verkhniy Urup isolated terrain (Karachay-Cherkessia) and called “Kazbich” was sent to the Moscow Zoo by Grand Duke Mikhail Nickolaevich, the vicege- rent of the Caucasus, there his traces were lost, and nothing is known about his future. In the wonderful book “The Animals of Russia” (Turkin, Satunin, 1902), the authors expressed doubts about that fact. However, Dinnik wrote (1910, p. 142): “…a calf was really delivered and placed to the Moscow Zoo. I know it from hearsay of S.A. Usov, my teacher of zoology. It is known that Professor Usov himself asked Grand Duke Mikhail Nickolaevich to send a Caucasian wi- sent as a present to the Moscow Zoo, and his request was satisfi ed”. One more young bull called “Kazan” was captured in 1899 at the Kisha River, and it was brought to the Bialowieza Zoo to get progenies from Lowland European bison to renew the generation and avoid inbreeding. About “Kazan” we know that it lived for a long time in one open-air cage with other animals but after 1903 its traces were lost. Whether it left offspring that could have been driven away from the Bialowieza Forest and could have survived is unknown though, of course, it was possible. A most interesting bull in the history of the Caucasian wisents is a bull called “Caucasus” that was captured there in 1907 and also brought to the Bialowieza Forest. In a year the Russian Emperor presented that wisent to Hagenbeck from Hamburg, a natural scientist and organizer of zoos and gardens, where “Cauca- sus” lived till 1922. After that it got to Count Arnim and was transported from Chapter 2. The Wisent in the Caucasus 37 Hamburg to Boitzenburg where it died three years later on February 26, 1925, but it had managed to leave seven calves from the cows of the Lowland line. Therefore, only one bull (“Caucasus”) and its seven progenies afterwards be- came ancestors of the new wisent line — Lowland-Caucasian (Bison bonasus bonasus x Bison bonasus caucasicus). Catastrophic general condition of the population was discovered as a result of the fi rst wisent census performed in 1926, based on which it was found out that only 52 animals had been preserved in the world! With that those animals were preserved only in the zoos and parks mainly in the West-European countries. Great attention drawn to that problem and objective approach to solving the tasks of the species preservation gave positive results, but growth of the popu- lation was very complicated and slow, and it was explained by life in captivity where they poorly bred. Nevertheless, as of January 1, 1938 there were already 96 European bison in the world, out of which 35 species were kept in Germany, 30 — in Poland, England and Switzerland — nine species in each, eight animals were in Holland, four ones – in other countries, and one European bison called “Bodo” dwelled in the territory of the USSR in the Askania-Nova Nature Re- serve (Jurgenson, 1956). It was the great grandchild of the famous “Caucasus” from Boitzenburg, it was bought from Rue company and brought to the USSR in September 1933 when it was three years old. Restitution of thoroughbred moun- tain wisents in Russia started with “Bodo”. In such countries as Poland, Germany and Sweden special programs for wi- sent breeding were developed, and in 1944 in spite of the 2nd World War the total amount of the European bison reached 146, but by the end of the war their quantity decreased again down to the level of 1937. As of January 1, 1947 there were 93 thoroughbred European bison in the world, the majority of which (44 animals) were preserved in Poland, in the western part of Germany (17 animals), in Sweden (16 animals), in the USSR (six animals), in England and Holland (four animals in each). The main work aimed at restitution of thoroughbred wisents in the territory of the former USSR was done in the state nature reserve called “Bialowieza For- est” where progenies of “Caucasus” were taken to in order to create a Lowland- Caucasian wisents line. During nine years starting from July 1946, 19 European bison were brought from Poland to the USSR, out of which 10 wisents — to the “Bialowieza Forest”, seven wisents — to the Central Wisent Nursery, and two wisents — to the Caucasian Nature Reserve. Careful work performed by the scientists drew the idea of the wisent restitution in the country nearer and nearer. However, it was possible to speak quite conditionally of the restitution of the Caucasian race because as a result of the accumulation cross breeding, in the mid-50s animals already had an insignifi cant admixture of “Caucasian blood”. Therefore, the thoroughbred Lowland-Caucasian wisents descending from the 38 U.A. Semenov The Wisents of Karachay-Cherkessia progenies of “Caucasus” are in their essence interracial hybrids. Nevertheless, they had distinctive characters; shorter curly coat especially on the head, large size, and strongly developed skeletal muscles. As for the last two characters there was an opinion that it was a result of inheritance of the hybrid vigor (het- erosis), as a result of the intraspecifi c hybridization in case of positive infl uence of choosing for cows bred in captivity a wild bull which is not their relative (Jurgenson, 1956). To develop the idea of the wisent restitution, Central Wisent Nursery was established on the basis of Prioksko-Terrasniy State Nature Reserve. The fi rst two species, “Puslav” and “Pustulechka”, were brought there from Poland on November 2, 1948. The second wisent nursery was established on the basis of the Okskiy State Nature Reserve in 1959. During three years, from 1959 to 1962, 11 animals were brought there from the Prioksko-Terrasniy Nature Reserve, and in 1967 and 1974 some more wisents were brought from the Bialowieza Forest. Work aimed at restoration of the wisents defi ned several major lines: thor- oughbred Lowland line, Lowland-Caucasian line of the interracial hybrids of thoroughbred wisents and the hybrid line of European bison and American steppe bison. A separate area was cross breeding of different combinations of the European bison, bison, and American-European bison hybrids with domestic cattle though this area of activity did not go much further. The next stage in the European bison history was restitution of the species within its natural and historical habitat in the Caucasus. For the fi rst time such activities were started in the Caucasian Nature Reserve where fi ve hybrids (American-European bison) were brought to in 1940, 15 thoroughbred wisents of the Lowland-Caucasian line were additionally brought there from the Bialow- ieza Forest, Prioksko-Terrasniy and Khoperskiy Nature Reserves in the period from 1948 to 1957. Later on the thoroughbred Lowland-Caucasian line (Bison bonasus bonasus x Bison bonasus caucasicus) was brought in to the North Os- setia, the Chechen-Ingush Republic and Karachay-Cherkessia. Chapter 3. History of Release and the Population Dynamics of the Wisent in the Teberda Nature Reserve

As of January 1, 1951 according to the State Studbook of Wisents and Bison (1956), in the USSR territory there were 83 animals, out of which 22 were thor- oughbred European bison, 12 were bison and 46 were hybrid European bison of different generation of accumulation cross breeding. In the Caucasus at that time there were hybrid forms of wisents and bison in the territory of the Caucasian Na- ture Reserve in the amount of 27 species and one thoroughbred European bison. In 1952 the second wisent park was organized in the Caucasian Nature Reserve (the fi rst one was created in 1940 at the Kisha River) at the Malaya Laba River (the isolated terrain of Umpyr), where two more young stud bulls called “Belyak” and “Berkut” born in the Bialowieza Forest were brought to, and the Central Wi- sent Nursery provided a one-year-old bull called “Murash” to the Umpyr Wisent Park. As a result, the Caucasian Nature Reserve received 22 thoroughbred stud bulls (Lowland-Caucasian line) in addition to their main herd of wisent and bison hybrids in order to restitute the wisent population in the Caucasus and perform ac- cumulation cross breeding in the wisent parks (Nemtsev et al., 2003). While defi ning the places for the wisents’ release the main focus was on pro- tected nature territories but the releases were the most successful in the nature reserves. This was explained by several factors: the protection there was better organized, and that was especially noted during the collapse of the USSR, scien- tifi c research was performed under control of highly-qualifi ed researchers, and that allowed to monitor the population, collect data about the ecology and biol- ogy of animals placed in the new environmental conditions, and fi nally quickly make decisions related to the wisent biotechnology and protection in case the animals left the protected nature territories. One of the most important factors of wisents’ surviving in those or other places was the availability of the staff re- searchers in the organizations where the wisents had been released. Writing the articles they often contained the increasing pressure of poachers’ hunting draw- ing the attention of the organizations to the problems arisen at the local level. A large-scale program of the wisent restitution within its historical habitat cov- ered signifi cant areas in the North Caucasus: 35 animals were brought to the Na- lchik Hunting Reserve of the Kabardino-Balkaria from 1959 to 1967; 48 animals 40 U.A. Semenov The Wisents of Karachay-Cherkessia were brought to the Tseyskiy Nature Reserve and the North Ossetian Nature Re- serve of the Republic of North Ossetia from 1964 to 1968. In 1987 the wisents also appeared in the Sunzhenskoe Hunting Reserve in the amount of 10 species, and in 1970–1975 in the Assinskiy Nature Reserve in the total amount of 49 species. In 1969 they were brought to the Ismaillinskiy Nature Reserve of Azerbaijan. In Karachay-Cherkessia the wisents appeared again in autumn of 1968 (Fig. 6). The decision was taken at the 3rd All-Union Meeting on the problem of the wisent restitu- tion in the USSR, and it was approved by the Action Plan of the Directorate General for Nature Protection, Nature Reserves and Hunting Reserves of the Ministry of Ag- riculture of the USSR No. 233-yu dated May 12, 1968. The fi rst eight animals among which there were two cows and six bulls arrived at the Kizgych Forestry, the Arkhyz part of the Teberda Nature Reserve, on September 5, 1968 (Table 1).

Figure 6. The Certifi cate of Delivery of Eight Wisents from the Okskiy Nature Reserve to the Teberda Nature Reserve dated September 5, 1968. From Archive Records of the Okskiy State Nature Reserve. Chapter 3. Wisent in the Teberda Nature Reserve 41 Table 1 The List of the Wisents Born in the Wisent Nursery of the Okskiy State Nature Reserve and Delivered to the Arkhyz Part of the Teberda Nature Reserve in 1968 (According to archive records of the Okskiy Nature Reserve (ONR))

Parents No. Date of No. Gender Name ONR/ Birth No. No. RZR Father Mother RZR RZR

September 1 Male Mechtatel 39/1913 Murmansk 1241 Musha 1316 13, 1965

June 01, 2 Male Merovey 42/2027 Muromets 1043 Mukha 1234 1966

July 09, 3 Male Metsenat 43/2028 Murmansk 1241 Mukholovka 1236 1966

July 16, 4. Male Mezon 53/2163 Muromets 1043 Muzeyanki 1169 1967

June 28, 5 Male Meredit 55/2165 Murmansk 1241 Murzinka 1309 1967

September 6 Male Mednik 58/2168 Muromets 1043 Murmanka 1310 09, 1967

May 31, 7 Female Mel 49/2159 Muromets 1043 Meduza 1504 1967

June 02, 8 Female Mecha 50/2160 Murmansk 1243 Musha 1316 1967

The second group of animals was brought from the Prioksko-Terrasniy Nature Reserve on November 3 of the same year, and there were two one-year-old bulls and six cows born in 1966 and 1967 (Table 2). 42 U.A. Semenov The Wisents of Karachay-Cherkessia Table 2 The List of the Wisents Born in the Central Wisent Nursery of the Prioksko-Terrasniy Nature Reserve (PTNR) and Delivered to the Arkhyz Part of the Teberda Nature Reserve in 1968 (According to the Studbook of the Central Wisent Nursery of PTNR)

Parents Date of No. No. Gender Name Birth RZR No. No. Father Mother RZR RZR

October 16, 1 Female Muya 2045 Beg 872 Muravka 801 1966

December 2 Female Murga 2046 Beg 872 Pustulechka 696 04, 1966

June 17, 3 Female Muska 2174 Beg 872 Mutovka 1317 1967

July 08, 4 Female Mussala 2175 Mushir 1608 Muskva 1389 1967

July 29, 5 Female Munka 2176 Mushir 1608 Mufta 1024 1967

August 29, 6 Female Mulyanka 2179 Mushir 1608 Murka 951 1967

August 05, 7 Male Mudrets 2177 Beg 872 Mushka 952 1967

October 06, 8 Male Mulat 2180 Mushir 1608 Murashka 1089 1967 Chapter 3. Wisent in the Teberda Nature Reserve 43 Table 3 The List of the Wisents Born in the Wisent Nursery of the Okskiy State Nature Reserve and Delivered to the Arkhyz Part of the Teberda Nature Reserve in 1978 (According to archive records of the Okskiy Nature Reserve (ONR))

No. Parents Date of No. Gender Name ONR/ No. No. Birth Father Mother RZR RZR RZR May 06, 1 Male Mervin 166/3817 Belkin 2021 Mednaya 1622 1976 July 10, 2 Female Mera 171/3822 Meteor 1624 Murzinka 1309 1976 July 09, 3 Male Melegar 169/3820 Mushan 2906 Muzeyanka 1169 1976 October 01, 4 Female Melnitsa 174/3825 Belkin 2021 Melfa 2025 1976 May 02, 5 Female Meliniya 164/3815 Belkin 2021 Medira 2032 1976 July 10, 6 Male Metall 170/3821 Belkin 2021 Metel 1614 1976 September 7 Male Metsamor 172/3823 Belkin 2021 Mechta 1621 14, 1976

A bull and a cow out of the brought wisents were left in the open-air cages of the Teberda Nature Reserve to be demonstrated to the visitors (Table 4), the remaining 14 animals without temporary holding were released to the territory of the Kizgych Chine (Bobyr, 1999).

Table 4 The List of the Wisents Born in the Central Wisent Nursery of the Prioksko-Terrasniy Nature Reserve (PTNR) and Brought to the Open-air Cages for Demonstration to the Visitors (According to the Studbook of the Central wisent nursery of PTNR)

Родители Date of No. No. Gender Name No. No. Date Birth RZR Father Mother RZR RZR October 1 Male Muster 07.06.2000 9464 Moris 0985 Mukharka 8621 16, 2003 October 2 Male Urbi 13.06.2001 2046 Riziko 7164 Luta 6634 16, 2003 December 3 Female Mubtika 18.05.2006 10775 Shporu 8841 Uriella 9451 12, 2007 44 U.A. Semenov The Wisents of Karachay-Cherkessia The Arkhyz part of the Teberda Nature Reserve, the Kizgych Forestry, is a chine elongated for over 20 km along the mountain river of Kizgych1, starting from the surroundings of the village of Arkhyz and limited on two sides by the rocky moun- tain ranges of Uzhum — 3,126 m above the sea level, and Cheget-chat — 2,963 m above the sea level, adjoining the spurs of An-Chatyr mountain — 3,434 m above the sea level, Sofi a mountain — 3,637 m above the sea level, Chuchkhur-Bashi mountain — 3,510 m above the sea level, and Psysh mountain — 3,489 m above the sea level, correspondingly, and coinciding with the borders of the Russian Fed- eration along the Greater Caucasus Mountain Range in the south-eastern, the high- est part of the nature reserve. According to the results of the latest forest surveying carried out in 2005, the total area of the forestry amounts to 19,272 ha (as a result of adjustments it decreased by 7 ha). Relief of the valley of Kizgych is a high-mountain relief with a slightly slop- ing and in some places wide (up to 600–800 m) fl oodplain which is gradually narrowing while moving up the stream, and having steep slopes on each side. A lot of small creeks fl ow into the river from both sides, and step by step they form a strong current in the middle and lower parts of the river. The lowest point of the valley near the river mouth is situated at about 1,500 m above the sea level. The dominant species in the forest stand of the fl oodplain forests are birch, grey and black alder, aspen, maple, hornbeam, etc. While moving towards the spurs of the mountain ranges and farther on to the slopes in the forests, there appear such species as oak, beech, elm, maple, gradually giving way to Nordmann fi r, eastern spruce, and scots pine. On the slopes of the mountain ranges on both sides of the river over 30 big and small avalanches slide down every year form- ing permanent cones in some places with the channel width of up to 100 meters and more in the lower part. Total area of forest lands of the Kizgych Forestry amounts to 8,813 ha (Table 5), the area of lands covered with forests is 8,717 ha. All of them are classifi ed as group 1 forests. In the forest reserve the prevailing plantations belong to the Nordmann fi r — 2,854 ha (32%), birch — 2,547 ha (29.2%), and pine — 2,183 ha (25%). Lands that are not covered with forests with the total area of 95.6 ha are represented by natural open forests — 51.9 ha, burnt places — 23.7 ha, and grassy clearings — 20 ha.

1 The Kizgych River rises from the foot of the northern slopes of the Greater Caucasus Moun- tain Range near Kizgych-Bashi mountain, where the riverbed is very narrow and runs at the bottom of the narrow rocky canyons. It is the right tributary of the Bolshoy Zelenchuk River which it fl ows into near Arkhyz. “K’yskhych” or “Kyskhach” are translated from the Kara- chay language as a “vice” or “gripe”. The word is based on “k’ysyk” — “narrow” or “com- pressed”, “ich” — “internal surface” or “part”. With the time the word transformed into the “k’ysygy’ych” fi rst (Tverdiy, 2011), and then into the “Kyzgych” and “Kizgych”. Author’s note. Chapter 3. Wisent in the Teberda Nature Reserve 45

Figure 7. The Kisgych Chine. Photo by U. Semenov

Figure 8. Dynamics of the Calves’ Birth.

The main part of non-forest lands (54% of the total forestry area) is covered by the alpine and subalpine meadows — 3,762.3 ha, which consist of the subal- pine tall grasses with frequent occurrence of rocky outcrops, the subalpine reed geranium-hedysarum meadows and the alpine barren lands. Other categories of 46 U.A. Semenov The Wisents of Karachay-Cherkessia non-forest lands also account for a signifi cant area: rock debris — 3,077.4 ha, barren rocky slopes — 1,729.6 ha, and glaciers — 1,486 ha. In spite of quite a large territory of the forestry not all its lands can be used by the wisents. Such land categories as the rocky areas, rock debris, swamps, gla- ciers, rockfalls, screes, bench gravel and landslides taking about 6,337 ha, which is approximately 33% of the area, are not used by them. A signifi cant specifi c weight falls within the alpine meadows — over 3,762 ha, about 50% of which are also inaccessible for the wisents (Table 5).

Table 5 The Land Area of the Kizgych Forestry of the Teberda Nature Reserve by Land Categories (According to the Forest Surveying of 2005)

Land Category Area, ha Natural Plantations 8,707.5 Plantations with Artifi cially Regenerated Species 1.8 Forestry Crops 7.7 Natural Open Forests 51.9 Burnt Places 23.7 Grassy Clearing 20 Hayfi elds 155.1 Meadows 3,762.3 Lakes 20.3 Rivers 76.6 Creeks 19.2 Dead Channels 1.6 Earth Roads 8.9 Paths 5.4 Other Clearings 0.8 Forestry Offi ce 0.5 Steep Slopes 1.9 Barren Rocky Slopes 1,729.6 Rock Debris 3,077.4 Swamps 2.2 Glaciers 1,486 Rockfalls 0.2 Screes 13.5 Landslides 12.6 Bench Gravel 15.5 Avalanche Cones 69.8 Total 19,272 Chapter 3. Wisent in the Teberda Nature Reserve 47

Figure 9. The Wide Glades at the Floodplain Terrace together with Clumps of Mixed Stand Cre- ate Favourable Feeding Stations for the Wisents throughout the Year. Photo by U. Semenov.

Figure 10. The Floodplain of the Kizgych River. Photo by U. Semenov. 48 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 11. The Forest Glades, the Most Important Feeding Stations of the Wisents. Photo by U. Semenov.

Thus, approximately 8,218 ha (or 42%) of the Kizgych Forestry area is practi- cally not visited and used by the wisents. These land categories, enriching the variety of the mountain ecosystems’ biotopes, in reality complicate and limit the wisent’s habitat and often become impassable barriers in their way to the lands suitable for seasonal locations of these animals. During the fi rst three years the wisent’s population did not increase because there were no adult cows in the groups. The fi rst calf appeared only in 1971 (Tara- sov, 1975). At that moment there were fi ve sexually mature females in the herd (two females out of the brought animals had died). It was left undiscovered what the nature of rutting was, as well as the animals’ activity rate and the number of animals participating in it, how many cows were in heat, how many cows mated bulls, whether the calves born by other cows died, or they did not have any calves that year at all. The next year the situation was the same, but as no dead calves of the current year were found, it could be assumed that in 1971 and in 1972 there was only one calf born in each year. In 1973 the observers noted that two calves were born, and the total number of animals reached 16. Birth of two females, and a male and a female after that, almost equalized the sex ratio in the population after death of two females resulting in 1:1 ratio. Two calves (females) were born in 1974, and the total number of animals reached 18. The next year was more productive, and three calves were born in the herd. Three out of eight adult cows had calves that year, and that amounted to 38% of the total number of sexually mature cows in the population. In spite of the Chapter 3. Wisent in the Teberda Nature Reserve 49

Figure 12. The Wisent Population Dynamics in the Teberda Nature Reserve. calves’ birth the population did not increase because two bulls and one cow died in 1975/1976. Only one calf of the current year was found in the herd in 1976. At that time the number of cows exceeded the number of bulls, and the sex ratio was 1:1.4, correspondingly. Nine out of ten adult cows were left without calves, i. e. 90% of cows were left farrow. Total number of animals stayed at the level of 18 species. Three calves were born the next year, but because of the death of two bulls the number of animals increased only by one and reached 19 (Fig. 8). In spite of the birth of calves (from 1968 to 1977 the population increased by 9% at the average), the wisents population did not increase signifi cantly. It is known that the cows if they are in good condition can bring forth every year or once in two years. The adaptation of the wisents used to plain environment was diffi cult in the mountains. That is why the birth rate in the herd was low. The cows’ fertility within that period amounted to 10–38%, 19% at the average. Un- fortunately, it is not possible to defi ne whether all the brought cows gave birth to a calf, or whether during that period some species brought forth twice. The dif- fi culty of the wisent’s adaptation is proved by the fact that two adult cows died in 1969–1970, then one cow broke a foreleg and a hind leg while crossing the rock debris of a rocky area, and the other cow fell down from the snow cornice over the deep gully. The mountain conditions to which the animals were not ready were the cause of both death cases of the adult animals. For nine years of the wisents’ life in the Kizgych Valley of Arkhyz their popu- lation reached 19 animals, i.e. the total number of animals had increased by fi ve species only. At that time the brought animals started to approach to the critical point of their reproductive ability, 13–14 years. At the same time the age of the highest sexual activity of the bulls is seven to eleven years, though they continue 50 U.A. Semenov The Wisents of Karachay-Cherkessia to participate in the rutting until the age of 14–15 years. The cows can give birth to calves until the age of 12–14 years, after that their fertility decreases (Zablots- kiy, 1957; Korochkina, 1971; Belousova, 1986). Death of four more adult wisents during 1975–1977 is marked in the Nature Records of the Teberda Nature Reserve. Sex-age structure of the wisent popula- tion of the Kizgych Valley before the delivery of the new animals was as follows: three adult bulls, eight adult cows, three young bulls aged under three years, two young cows aged under three years, and three calves born that year, among which there was one female and two males. In spring of 1978 the third group of animals consisting of four bulls and three cows was brought to the valley of Kizgych (Table 3). This decision was made in order to speed up the rate of breeding and to avoid inbreeding among the earlier released wisents. Three calves were born that very year. By the time when the new wisents were released the number of animals in the herd had reached 22. As a result the total number of animals in summer 1978 was 29 (Fig. 12). Release of the new animals shifted the rutting season time and led to a certain disorganization of the animals within that period. Neverthe- less, the observers mentioned that two calves of the current year appeared in the herd in the beginning of summer 1979. Only two out of 13 sexually

Figure 13. The Kizgych Floodplain, the Middle Course (Frontal View at the Abishara-Akhuba Mountain Range) Photo by U. Semenov. Chapter 3. Wisent in the Teberda Nature Reserve 51

Figure 14. The Combination of Glades, the Floodplain Deciduous and Fir Forests Create a Vari- ety of Comfortable Biotopes for the Wisents’ Life in the Valley of Kizgych. Photo by U. Semenov.

Figure 15. View at Sofiyskoe Sedlo Mountain via Which the Wisents Often Go to the Neigh- bouring Isolated Terrain of Sofia. Photo by U. Semenov. 52 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 16. The Upper Part of the Kizgych Chine, the Place Where Wisents Get to the Isolated Terrain of Bugoy-chat. Photo by U. Semenov

mature cows delivered calves, the rate of farrow cows in the herd amounted to 75%. Three calves were born each year in 1980–1981, and four calves were born each year in 1982 and in 1983 (Bobyr, 1992). Thus, in spite of the fact that the animals left the Kizgych Valley for the adjoining territories, and the population decreased, the release of the new wisents had a positive effect, and the birth rate increased up to 14% in 1982. With the growth of the total number of the animals, and periodical f ormation of new groups of different size and sex-age structure, and also constant migra- tion of the animals within the chine territory, within the nature reserve territory and beyond its borders, it became rather diffi cult to perform accurate registration of the wisents. To determine the total number of the wisents and the population sex-age structure recording was performed three times in 1982. The fi rst stage was performed during March 12–15. By that time a lot of snow had fallen in the mountains, and the wisents moved towards the lower part of the river valley, and stayed in the fl oodplain terraces feeding on dead grass in the open areas of de- ciduous species and shrubs. The main principle of recording was as follows: the recorders came to the observation stations, observed the surrounding territories through the fi eld glasses and registered all the discovered animals. In the places where visual examination was hindered the observers went along the routes and discovered the wisents by their traces. In such a way 23 animals were discovered Chapter 3. Wisent in the Teberda Nature Reserve 53

Figure 17. Extensive Avalanche Cones are Important Feeding Stations of the Wisents in Spring- time. Photo by U. Semenov. in the territory of the nature reserve, and six animals were registered beyond its borders, in the isolated terrains of Shorovo and Linevaya Balka. In May of the same year the Science Department of the nature reserve together with security offi cers carried out the second recording by means of registration of the animals’ occurrence in the 22 km long linear route along the Kizgych fl oodplain. Based on the recording results the total number of the wisents was 31 animals taking into account four animals which had returned from the Linevaya Balka and two animals staying in the isolated terrain of Shorovo. The third stage of the recording work took place in autumn, on November 17–19. By that time a 10–15 cm snow cover had formed, but on the slopes of the southern expositions and in the fi r forests there was almost no snow. At that time the observers divided into two groups in order to cover the biggest part of chine territory at a time. The fi rst group examined the fl oodplain of the river and the adjacent slopes up the stream from the Perviy Post through to the Bugoy-chat isolated terrain, where they found 12 wisents. The second group of the observers explored the fl oodplain and the surrounding territories down the stream through to the suspension bridge. They found traces of a group of wisents and two soli- tary animals. 54 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 18. Avalanche Cones Overgrowing with Young Vegetation Become One of the Most Attractive Feeding Stations. Photo by U. Semenov Chapter 3. Wisent in the Teberda Nature Reserve 55

Figure 19. Headstreams of the Kizgych. The Highest Part of the Chine is not Visited and Used by the Wisents. Photo by U. Semenov.

On the next day all the recorders were sent to that area to fi nd out the quan- tity of animals in the group, and they managed to discover a small mixed herd consisting of 10 animals, and a group consisting of two adult bulls. During the examination near the Grushevaya Polyana the recorders saw one more group consisting of three species which kept apart from the earlier discovered animals. Twenty-seven wisents were registered in the territory of the nature reserve as a result of the autumn recording. However, as some observers said, the total amount of the animals was approximately 40 wisents. It was thought that the existence of the undiscovered groups consisting of eight to fi fteen animals was possible, but as it was rightly mentioned in the re- port, it needed clarifi cation. The population decrease was explained by the death of two wisents and the animals’ leaving the nature reserve, though the observers mentioned that there were calves of the current year in the groups. For the fi rst time from the moment of the species reintroduction the bulls prevailed in the sex ratio of the registered animals — 1.3:1, and the cows’ fertility reached 57%. Four out of seven adult cows found in the territory of the nature reserve had calves. Probably several cows which could have given birth to calves that year stayed beyond the nature reserve territory and were not registered, but there was no in- formation on occurrence of calves of the current year in the neighbouring lands. 56 U.A. Semenov The Wisents of Karachay-Cherkessia In 1983 the registration of the wisent population was carried out on March 13–16, after the snowfall, by means of visual counting of the animals near the hay storages where they came to feed. The specifi cation of data was made by tracking their traces and the registration of cases when the wisents were seen, while going along the route in the river valley. The total number was 33 animals with the species density amounting to 3.6 animals per 1,000 ha of the forest area. Birth of four calves was observed in the same year. In May of the next year the observers carried out an interim recording, as a re- sult of which they registered 37 animals of different gender and age, later on they registered the appearance of three newborn calves. Thus, the total animal number amounted to 40 with the species density of 4.6 animals per 1,000 ha. The sex ratio in the population was 1:1.2, where the number of cows exceeded the number of bulls. Two calves were born in 1985, and the total number of animals reached 44. It was explained by the appearance of two bulls in the valley which kept beyond the nature reserve territory before that. Only two out of 18 sexually mature cows had calves. The rate of farrow cows in the groups amounted to 89%. However, the actual number of the cows participating in the rutting and giving birth to calves was probably a bit higher because the analysis of the population dynamics showed that in spite of the registered number of the calves of the current year, the population was increasing faster. Probably, it is explained by the underes- timation of the calves’ quantity during the parturition years, and adding them to the total number of animals as young or adult ones during the recording and registration. It happened during the regular recording of the population in Janu- ary 1986 when the observers registered 51 wisent and only three calves of the current year. A similar winter recording, which was carried out one year later, showed rather poor growth by only three animals (54 animals), in spite of the record number of born calves (six animals). The reason for such poor increase of the population was a severe and snowy winter. At that time a lot of wild un- gulates died of exhaustion and in the avalanches in the Teberda Nature Reserve. During the examination of the lands they discovered the fragments of carcass, fur, bones and skulls of 44 turs, 10 wild boars and four wisents, three of which were females and the gender of one wisent was not defi ned. It is worth mentioning that the winter of 1986/1987 was snowy and favourable for avalanching. While the turs mostly died in the avalanches, other ungulates, including the wisents and wild boars became exhausted from moving in deep snow. It was impossible to restore the expended energy because it was diffi cult to fi nd food, and that led to their weakening and death of exhaustion. When the observers examined the dead wisents they discovered that they didn’t have depot fat at all, and in their stomachs there was only dead grass. In 1988 the wisent population decreased dramatically down to 39 animals. The winter of that year was less snowy than that of the previous year, and only Chapter 3. Wisent in the Teberda Nature Reserve 57 one wisent which had died of old age was found. In spite of a relatively mild winter of 1987/1988 the nature reserve staff found wild boars which had died of exhaustion near the Stariy Kordon, and not far from the Central Manor. The main part of the wisent population stayed in the Kizgych Valley but during sum- mer and autumn it was observed that solitary animals and several groups consist- ing of three to six animals went out to the area of the Krivaya Balka and down the stream of the right bank of the Bolshoy Zelenchuk. Probably some wisents did not return, and nothing was known of their further fate. Certain increase of the population was caused by the birth of four calves, after which the number of animals increased up to 43. The next year four more calves were born, and the population increased up to 47 animals with the species density of 5.2 animals per 1,000 ha. Some groups which had migrated to the neighbouring lands were not in the scope of the recording. According to the expert opinion of the observ- ers, within that period two groups consisting of four and six animals each (about 10 animals in total) were outside the nature reserve territory. The registration of the number of animals during the subsequent years confi rmed that assumption. Solitary animals and small groups were periodically found outside the territory of the nature reserve. In 1991 the animal counting performed in several stages allowed to cover the whole chine quite fully. Taking into account the wisents

Figure 20. The Steep Rocky Slopes, Deep Cracks and Thick Birch Crook-Stem Forests Restrict the Wisents’ Movements in the Kizgych Chine. Photo of U. Semenov 58 U.A. Semenov The Wisents of Karachay-Cherkessia which lived outside the nature reserve territory, in the surroundings of the Zelen- chuk Observatory, the total number of animals amounted to 55. In the beginning of the 1990s the eastern border of the wisents’ habitat lay from the headstreams of the Bugoy-chat originating from the confl uence of creeks on the slopes of the Greater Caucasus Mountain Range, and father along the Uzhum Mountain Range, the western border started from the confl uence of the Kizgych River and the Bugoy-chat River and ran along the slopes of the dividing ridge up to the confl uence of the Psyzh and the Sofi a. To the north of the source of the Bolshoy Zelenchuk River and up to the village of Nizhniy Arkhyz where several wisents had lived before that (Bobyr, 1989), the wisents were not observed. The security offi cers of the nature reserve said that the residents of Arzhyz had seen wisents near the Tserkovnaya Polyana situated at the fl oodplain terrace of the left bank of the Psyzh River. That meant that the wisents’ range had decreased signifi cantly. Earlier the total territory of their habitat had been approximately 50 thousand ha but at that time it did not exceed 25–28 thousand ha. The total popu- lation decreased by eight animals and amounted to 47. The majority of animals was registered along the Kizgych where the population density was fi ve animals per 1,000 ha of the forest area. The main reason for the population decrease dur- ing that year was the death of eight animals caused by exhaustion during moving in deep snow in snowy winter of 1991–1992. The analysis of animals’ population performed in 1993 based on the recording performed on April 5 showed that there were 48 animals taking into account one dead wisent and two newborn calves. The next year the population recording was not carried out in Arkhyz, but the observers saw two calves of the current year in the herd, and as a result the number of animals amounted to 50. Three calves of the current year were registered in the groups in 1995. The fi rst group consisting of 14 animals was found near the Byky isolated terrain, the second group consisting of 20 animals — near the Tomak-kaya, and the third group consisting of approximately 13–15 animals — near the confl uence of the Psyzh and the Kizgych. Thus the number of wisents was 47–49. It was supposed that a group of three to fi ve animals lived in the adjacent territories. It was evaluated that there were 53 wisents. In 1994, 1995, 1996 the rate of farrow cows reached 94%. Only two or three out of 33 registered sexually mature cows had calves of the current year. The wisents started to go out of the nature reserve more often and stayed in the neighbouring lands for a long time not returning to the nature reserve for wintering. In 1996 due to the deep snow cover in was not possible to perform planned recordings covering the territory of the mountain valley and the adjacent lands. According to the information of the nature reserve staff, who observed the ani- mals during the period of their mating aggregations, in the river valley there were several herds consisting of 38 animals in total, and there were four calves Chapter 3. Wisent in the Teberda Nature Reserve 59

Figure 21. Melting Snowflakes and Glaciers Form a Lot of Creeks in the Territory of the Kizgych Chine, and in Summer that Allows the Wisents not to Go Down from the Subalpine Meadows to the River Valley for Watering. Photo by U. Semenov. 60 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 22. The Arkhyz Population of the Wisents as of May 5, 2012. Photo by U. Semenov

born that year among them. Presumably, about 15 animals belonging to different groups lived in the adjacent territories outside the nature reserve. The new re- cording in the beginning of the next year did not confi rm the existence of a large unregistered group. The total number of animals was 43, and it was defi ned by means of tracing their paths and visual examination. Only some animals were registered by visual examination. Forty-fi ve wisents were registered in 1998, and 46 animals were registered in 1999. The wisents started to leave the protected territory more often, and that did not allow the observers to perform a complete visual analysis of the whole wi- sent population. The method of wisent counting taking into account the groups possibly existing outside the nature reserve could not refl ect the real situation because there the wisents were not protected and observed by state inspectors. Those territories were not explored in order to fi nd animals which died for vari- ous reasons. In 2000 recording of the wisent population in the mountain valley of Kizgych was carried out in spring (in early April) under supervision of the author, by means of exploration of the wisents’ habitat with the simultaneous trace tracking in the places of their aggregation. The traces of a wisent group consisting of fi ve Chapter 3. Wisent in the Teberda Nature Reserve 61 species were found in the fl oodplain terrace and on the slopes of the right river bank near the waterfall (close to the Volchye Mesto isolated terrain), traces of a group consisting of about ten species were seen in the Byky isolated terrain, a herd of four adult bulls lived in the headstreams of the Kizgych River. The total number of the wisents discovered by traces (18 animals), and visually observed (two animals) amounted to 20. That year it was decided not to include the spe- cies possibly living in the adjacent territories into the number of the registered wisents. That decision was disputable from the point of view of certain employ- ees, but still that very number without changes was included into the Nature Records report for 2000, and the observations made in the subsequent years confi rmed the correctness of the taken decision and refl ected the real number of the wisents in Arkhyz at that moment. Appearance and subsequent disposition of the military forces in the headstreams of the Kizgych River in 1998–1999 aggra- vated the situation. Building of dugouts and blockhouses, usage of barbed wire for fencing, as well as fi res, fi ring exercises, systematic arrival of a helicopter landing in the river fl oodplain, big groups of people going to patrolling rounds, and many other things related to uncontrolled presence of the contract soldiers in the territory of the nature reserve drastically increased the wildlife distur- bance factor there. The camp organized by the soldiers was located in the path of seasonal migrations from the isolated terrain of Bugoy-chat to the spurs of the Sofi yskoe Sedlo Mountain Range where the wisents had gathered during mating aggregation periods before, in the same place there were natural and artifi cial salt sources. The animals were ousted from that important area. And the infl u- ence was so signifi cant, that even many years after the military people had left the place, only traces of the solitary wisents were seen there. The wisents had to leave the safest and the most comfortable places and stay in the areas where the disturbance factor was not so strong. Constant relocation of the military vehicles in the river fl oodplain, from the river source to the headstreams and back, left them no possibility to live safely in those conditions. The arrival of people and noisy military vehicles had a very negative impact on the wisents of the Arkhyz area of the nature reserve, which was an extended open river fl oodplain, having 500–800 m long fl oodplain terraces on both sides with the combination of glades and mixed forests giving way to coniferous forests going up to the high rocky mountains. Moreover, the arrival of contract soldiers brought not only wildlife disturbance, there were also poaching cases. According to the information from one of the soldiers, salted skins of wild animals and three wisent heads were loaded into the vehicle during one of the regular turnouts from the upper camp. In his opinion, the heads were prepared for making mounts. From 1996 to 2000 the observers found two wisent skulls in the avalanche cones of the Kizgych Valley; they found a lame and exhausted female wisent which probably died within a few days after it had been found, they also discov- 62 U.A. Semenov The Wisents of Karachay-Cherkessia ered the fragments of bones and the skin of an adult wisent killed by the wolves near the salt lick, and the wisent remains were found in the forest area (the true cause of its death was not defi ned). Presumably, the wisent was killed by the poachers. The prevailing conditions became critical for the population which was al- ready in the state of depression. It is very illustrative that only one calf was seen in the herds from 2000 to 2005! Even weakening of the anthropogenic pressure during the subsequent years could not change the situation, the number of ani- mals continued to decrease. The rate of farrow cows in the last ten years reached the critical level of 95–98%, аnd even 100% in certain years. The recording performed in 2004 showed that only 11 wisents lived within the nature reserve territory, and no wisents were met in the adjacent territories. Dur- ing 2005 recordings 15 wisents were found in the Kizgych Valley, among them there were seven adult bulls, one young bull and one adult cow, and a mixed group of six young and adult animals whose sex was not determined. The body fragments of an adult wisent having died in the avalanche were found in the ava- lanche cone in the same year. In 2006 the population amounted to 13 species. The examination of mineral springs’ outlets let them conclude that there was a calf in the mixed group, sex and age structure of which had not been defi ned. According to the Nature Records of the nature reserve, the wisent population ranged from 11 to 14 animals during 2006–2011, birth of calves was registered several times. However, the available data about that period cannot provide a precise picture for the detailed analysis. Application of rough estimates led to the erroneous picture of the Arkhyz population sex-age structure and the actual overestimation of the wisent number by almost three times. The wisent recording carried out by the author together with the nature re- serve staff during May 3–6, 2012, and the subsequent usage of hunting cameras for 180 days showed that only fi ve wisents lived in the territory of the Kizgysh Forestry of the Teberda Nature Reserve, among them there were two adult bulls, one old bull, one adult cow and one young bull aged about one year (Fig. 19). One of the bulls periodically left the group but then joined it again. The wisents ceased to leave the valley since 2009. According to the information collected in the adjacent territories, the wisents had not been seen there for several years. Chapter 4. Habitat Structure and Migration Dynamics

In general, the wisent may be deemed a sedentary animal; however, high mo- bility is characteristic of it in the conditions of mountain ecosystems both within a year and within a day. In the past, with more opportunities to migrate and lack of human interventions those migrations were associated with seasons. Depend- ing on the winter conditions, the animals went down to the lower territories where the snow cover was considerably thinner than in the high mountains, and then went back up the mountains to the upper border of forests following the retreating snow and the growing young greenery. In late June when almost no snow was left in the subalpine and alpine meadows, the animals stayed there for most of the feed time having rest in the shade of the nearby forest during the daytime. We know from the early sources that the native Caucasian wisent had clearly defi ned local seasonal migrations (Latsarius, 1879; Dinnik, 1884, 1910; Fila-

Figure 23. A Young Bull Aged Three to Four Years. Photo by U. Semenov. 64 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 24. An Adult Bull on the Forest Glade. Photo by U. Semenov.

tov, 1909; Bashkirov, 1939; and others). The animals went down to the foothills reaching Pregradnaya and Storozhevaya villages during the most severe winters with little snow. The other part of the animals stayed in the alpine zone with spacious meadow areas cleared from snow by the wind, so the animals were provided with food for the entire winter. It is possible that the seasonal migrations of the animals were of a larger scale in the 10–15th centuries when they could go to the mid-mountain, foothill and even steppe territories of the Caucasus but then their opportunities were lim- ited with the course of time and land reclamation. The increased population and easier chasing and hunting wisents in the plain territories of the region dramati- cally reduced the seasonal migrations of the animals making them focus on more localized territories in the high-mountain part of their habitat. In the 19th century, such migrations hardly exceeded 10 km, which was possibly due to the low den- sity of the species and the weak competition for localized areas more comfort- able during the winter period. The restored wisents also showed clearly defi ned seasonal migrations related to the snow cover almost in all the mountain territories. As noted by A.S. Nemt- sev (1988), large-scale migrations of the Caucasian Nature Reserve wisents to the foothills in autumn were observed once every 8 to 10 years. The length of such migrations oriented southeast to northwest reaches 30 km with elevation Chapter 4. Habitat Structure and Migration Dynamics 65 changes of around 2,000 m. According to other sources (Kalugin, 1958), such difference may reach 2,600 m. The scientists believe that the migration pro- cesses are not so clearly defi ned during the winters with little snow although the animals are constantly moving from station to station, the daily movement length hardly ever exceeds 1,000 m and sometimes even goes down to several hundreds of meters. As noted by V.E. Sokolov and A.K. Tembotov (2003), sea- sonal migrations of the wisents in the Malaya Laba River Valley were minimal, their length could be not exceeding one hundred meters. The altitudinal movement range of the Arkhyz population wisents is between 1,400 and 2,800 m above sea level but the animals come to high mountains during the summer time only. So the mountain populations of the wisents from the Tseyskoe Chine in North Ossetia, from the Arkhyz population in Karachay- Cherkessia and from the Caucasian Nature Reserve dwell in similar ecologi- cal conditions and at approximately same altitudes (Kalugin, 1977; Lipkovich, 1985; Bobyr, 1989, 1992; Semenov et al., 2001; Semenov, 2003). Wisents show sedentariness tendencies not only in their permanent habitation areas but also in the reacclimatization territories, moreover, the animals do not tend to explore the neighbouring territories even in overstocked populations. They go there more for the reason of young animals being ousted from their

Figure 25. A Group of Wisents on the Artificial Salt Lick. Photo by U. Semenov. 66 U.A. Semenov The Wisents of Karachay-Cherkessia main habitation areas. Such forced migrants sometimes go away for signifi cant distances. Some species covered 90 km in the Caucasian Nature Reserve, 160 km in the Khoperskiy Nature Reserve, 100 km in the Crimea away from their initial release place, and 40 km in Volyn and in the Tsuman Forest Reserve. Cases were reported when separate species covered 500–600 km (Zablotskiy, 1975; Kolisnyk, 1989; Pavlov, 1999; Nemtsev et al., 2003). The wisents that were brought to the Kizgych Valley in 1968 also had seasonal and daily migrations. For the fi rst time of their reacclimatization, such migra- tions were localized and chaotic as the territory exploration was in process. But in one year, in early August 1969, the whole herd covered around 20 km from their initial release place and moved to the neighbouring Psyzh Valley (Bobyr, 1992). Although the animals crossed the reserve borders from time to time (Ta- ble 6), they did not explore any new territories because of the weak growth of the population. As a rule, the animals that left the area returned after some time. The case when a bull migrated 8 km northeast from the river fl oodplain to the Chabakly Lake area in autumn 1972 may be deemed the fi rst attempt of oust- ing the animals from their main territory, but the wisent came back and joined the main herd as winter set in. In August next year, a bull (probably the same one) left the Kizgych Valley again, went down the Bolshoy Zelenchuk River to the Ermolovka village surroundings and even lower to Zelenchukskaya village where it was living for around one and a half year and then moved on to the foot- hill and steppe territories of Karachay-Cherkessia where the local people saw it in the surroundings of Adyge-Khabl village. That is around 150 km away from Arkhyz. Nothing is known about the further fate of that animal. It is noteworthy that such migrations of solitary bulls and even small groups happened mainly in the late summer and early autumn. As the rutting starts, and the competition grows abruptly, young species are ousted by stronger adult bulls and have to look for new territories and groups. A few years later, the number of migrations outside the Kizgych Valley in- creased. Groups with different gender and age structures were leaving the re- serve. The interdependence of the population growth and the number of migra- tions away from the initial release place was observed. There was not enough space for the wisents in the Kizgych Chine any longer. The observers noticed that the local migration processes activated, and the herd fell apart in 1973. The animals hardly ever gathered into one group. Even more active migration processes were noticed the following year, and they gath- ered only in winter when the snow cover reached the highest level. The wisents frequently grazed and had a rest on the subalpine and alpine meadows especially in the area of the Bugoy-chat isolated terrain, in the warm season. In winter, they kept to the mouth of the Kizgych River, on the Malaya and Bolshaya Che- gordala, and the Lebzor glades, and near Mertvoe Lake. Chapter 4. Habitat Structure and Migration Dynamics 67

Figure 26. An Adult Female Wisent in the Winter Forest. Photo by U. Semenov.

Several attempts of the animals to reach the left side of the Zelenchuk River were noticed. The wisents were crossing the river but could not go to the Abishi- ra-Akhuba Mountain Range slopes because of the wire fencing of the Arkhyz Forestry along the left side of the Bolshoy Zelenchuk and the Arkhyz Rivers. Those attempts to reach the slopes were probably due to their southern exposi- tion because the animals were trying to go there only in winter when the height of the snow cover in the Kirgyz isolated terrain is one meter and more. During the warm season, the wisents usually kept to the upper border of the forest and the subalpine meadows. In 1980, the same as during the previous years, the wisents used the Kizgych territory bordering blocks 7 and 21 in the north and blocks 81–88 in the south. Before that, the wisents had not entered the Chertova Melnitsa isolated terrain in the southern part of the reserve even in summer although the landscape and fl ora are similar there. While in the northern part of the Kizgych, they regularly passed to the Psyzh isolated terrain along their fi xed routes crossing blocks 19– 28 and 30. Two bulls kept in the area of the cattle farms in the Shorovo isolated terrain during the whole year. In summer, groups of animals consisting of 15 to 27 species several times showed up in the Sofi a isolated terrain that they reached crossing the Sofi yskoe Sedlo Pass. During the autumn periods of 1979 and 1980, the animals were more frequently seen at the upper border of the forest and on the subalpine meadows (blocks 73, 76–80, 82–83, 92). 68 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 27. An Old Wisent. Photo by U. Semenov. Chapter 4. Habitat Structure and Migration Dynamics 69

Figure 28. An Adult Bull Crossing a Wide Glade. A Floodplain Terrace of the Kizgych River Headwaters (May 4, 2012). Photo by U. Semenov. During the winter period of 1982–1983, the majority of the wisent population dwelled in the Kizgych, and two wisents only used to leave it for the Tserkovnaya Polyana territory visiting the timber cutting areas where they fed on needles and bark from the cut trees. Those wisents came back to the Revunok isolated terrain of the Kizgych Valley in spring, in the middle of March, and then went to the river headwaters. One bull migrated to the Psyzh isolated terrain in summer and moved on towards Pkhiya village through the mountain pass. Shepherds saw the wisent grazing 10 km away from the village near the path leading to the moun- tain pass in late July. The shepherds said that the wisent was then showing up in the area of the Temir-Kulak Pass in autumn, in the third decade of September. During 1986, the observers saw bulls going down the Bolshoy Zelenchuk River to the Linevaya Balka isolated terrain in spring, summer and autumn. One bull that was dwelling in the surroundings of the observatory returned to the Kizgych Chine but very soon went back again together with a young bull. Some bulls were noticed to leave the reserve for the Ginseng plantation area and the Zelenchuk observatory surroundings in the autumn period of 1987. In the late 80s, the border of habitation of the wisents released to the moun- tain valley lay from the side branches of the Uzhum Mountain Range through the Zelenchuk observatory surroundings and up the fl oodplain terraces of the Bol- 70 U.A. Semenov The Wisents of Karachay-Cherkessia shoy Zelenchuk River to the place of the confl uence with the Kizgych River. The western and north-western borders were limited by the slopes of the Sofi yskiy Mountain Range. In June 1989, the territory of the Karachayka River headwaters was examined, and traces of three wisents were found there, as well. According to the shepherds herding the cattle in the Pkhiya River headwaters, two adult wisents and one young species were seen there but they were not noticed in that place in the late summer any longer. Those wisents could possibly return to their former habitation area cross the mountain pass or could leave for the lower territories of the Bolshaya Laba. The wisents were said to be seen on the southern slope of the Greater Caucasus Mountain Range in the area of the Pskhu River headwaters but that information needed to be double-checked as G.Ya. Bobyr noted (1992). In spring, late April of 1989, some groups of wisents moved to the Kizgych River headwaters where they stayed for the whole summer and the beginning of autumn and then went downstream to the river mouth except the group consist- ing of nine species dwelling on the fl oodplain terraces in the middle river area. Migrations of some groups consisting of three to six animals, as well as of soli- tary animals were observed from midsummer till winter when all the animals came back to the reserve except for two animals that stayed in the observatory surroundings and two more ones that stayed in the Krivaya Balka. In the early 90s, the eastern border of the wisents’ habitat in the Kizgych Val- ley lay from Bugoy-chat Mountain along the Uzhum Mountain Range, the west- ern border – from the confl uence of the Kizgych and the Bugoy-chat Rivers and on along the slopes of the dividing ridge to the confl uence of the Psyzh and the Sofi a Rivers. The northern border lay from the origin of the Bolshoy Zelenchuk River to Nizhniy Arkhyz village where several animals dwelled in 1991 and 1992 but no wisents were observed there in 1993 any longer. They probably re- turned to their former dwelling. That year, the habitation borders slightly shifted to the north down the Bolshoy Zelenchuk River and covered the Linevaya Balka isolated terrain and the ginseng plantation surroundings where four adult wisents dwelled during the whole year. As you can see from the table, the wisents left the reserve quite frequently, and people reported seeing them almost every year. Some years, several groups and solitary animals stayed in the adjoining areas at the same time. The further reduction of the population by 2000 led to changes in the territorial migration activity of the wisents in the Kizgych Valley. The number of reported observa- tions dropped considerably although the animals were noticed outside the re- serve almost every year (Sofi a, Krivaya Balka, Linevaya Balka and other isolat- ed terrains). In August 2008, one mixed group of four wisents only was noticed (gender and age structure undetermined) in the Sofi a isolated terrain. Then the animals came back to the Kizgych headstreams area through the Sofi yskoe Sedlo Pass in the fi rst decade of September. Chapter 4. Habitat Structure and Migration Dynamics 71 Table 6 Table N/a in 5 days Outcome Kept in the Zelenchuk River probably came back Stayed there and then Went downstream the Went Zelenchukskaya village Came back the same way surroundings for 1.5 years km Covered Distance Approx. 150 Approx. 45 km Approx. 15 km Came back Approx. 15 km Came back in September Approx. 15 km Kizgych Valley of the Teberda Reserve Teberda of the Valley Kizgych

in 1975 Location many times. surroundings yskoe Sedlo Pass) yskoe Sedlo Pass) yskoe Sedlo Pass) fi fi fi Psyzh isolated terrain 20 km Driven back So So So Zelenchukskaya village a isolated terrain (crossed the a isolated terrain (crossed the a isolated terrain (crossed the fi fi fi continued to walk round the The adult bull that left for the Psyzh isolated terrain in early winter was not found although The wisent that left the territory Ermolovka village surroundings Approx. 20 km Noticed in Karachay-Cherkessia So So So Khabl village in September 1975. wisent traces were observed there foothills. Last report from Adyge- foothills. Last report from foothills near Adyge-Khabl village foothills near N/a bull) animals) Structure Mixed herd (all Gender and Age and Gender (supposedly, adult (supposedly, - Mixed herd - Mixed herd 1 Adult bull1 Chabakly Lake area Approx. 8 km Came back in early winter 8 Mixed group 12 Wisents Number of Number Description of the Wisents’ Migrations outside the Wisents’ Description of the 1976 1 Adult bull Psyzh isolated terrain Approx. 20 km 1977 Same bull 1972 1973 1971 1970 1969 Years (August) (August) (August) (August) September (September) 72 U.A. Semenov The Wisents of Karachay-Cherkessia ) continuation back there came back Came back cattle sometimes Spent winter there Came back by winter Table 6. ( Table mouth to spend winter N/a. Supposedly, came N/a. Supposedly, together with domestic and autumn there, came back to the Kizgych and Spent the whole summer joined the herd by winter Moved to the Psyzh River Came back as winter set in along the path. Supposedly, along the path. Supposedly, Noticed when moving back Stayed for one year grazing 5-20 km Came back Approx. 8 km Approx. 7 km Approx. 35 km Approx. 15 km Approx. 20 km Range terrains a and Arkhyz isolated a and fi a isolated terrain fi village surroundings So Psyzh isolated terrain Approx. 20 km Came back Arkhyz isolated terrain Arkhyz isolated terrain Arkhyz isolated terrain Approx. 20 km Shorovo isolated terrain observatory surroundings Tserkovnaya Polyana areaTserkovnaya 2 km Came back in March Zelenchuk River right side Linevaya Balka isolated terrain Psyzh, So Kept to the cut out strip along The wisents went downstream the Zelenchuk River right side, slopes of the Abishara-Akhuba Mountain of the electric power line in the Zelenchuk animals Adult bull Psyzh isolated terrain – Pkhiya Adult bull Adult bull Adult bull Adult bull Young bull Young Adult bulls Adult bulls different age different Adult bulls of Mixed groups Whole herd by and young bull different groups different Young bull out of Young the newly brought 2 1 2 4 2 1 2 29 2-13 July 1983 1983 1982 1980 1 1986 1979 1978 (winter) (March) (summer) (December) 2 N/a Chapter 4. Habitat Structure and Migration Dynamics 73 ) continuation left N/a winter winter winter winter winter then unknown herd by winter the Kizgych by winter Table 6. ( Table Revunok and Kazachya the Kizgych and joined Polyana isolated terrains as Spent the whole year there, autumn there, came back to autumn there, came back to Returned to the Kizgych by Returned to the Kizgych by Returned to the Kizgych by Returned to the Kizgych by Returned to the Kizgych by the observatory as cattle Moving to the neighbouring spring set in and returning to Spent the whole summer and Spent the whole summer and km km Approx. 12-15 Approx. 20-30 Approx. 25 km Balka isolated terrain Verkhnyaya Ermolovka village Verkhnyaya the Bolshoy Zelenchuk River to Went downstream the right side of Went Ginseng plantation area - Linevaya Zelenchuk observatory surroundings Approx. 10 km Zelenchuk observatory surroundings Approx. 10 km - Zelenchuk observatory surroundings age different agedifferent Group of females and young species of Bull group of different 221 N/a2 N/a1 Adult bull Zelenchuk observatory surroundings Adult bulls2 Approx. 10 km Adult bull1 Polyana area Tserkovnaya Spent winter there Krivaya Balka area Adult bulls Kazachya Polyana area Approx. 7 km Adult bull Ginseng plantation area Approx. 10 km Approx. 9 km Spent winter there Zelenchuk observatory surroundings Approx. 10 km 24 Adult bulls N/a13 Zelenchuk observatory surroundings Solitary bull Approx. 20 km 6 Zelenchuk observatory surroundings Approx. 10 km 1990 N/a 1991 1987 1988 N/a 1989 (January-April) 74 U.A. Semenov The Wisents of Karachay-Cherkessia ) end Table 6. ( Table N/a Came back All the animals came back All the animals came back km Approx. 9 km N/a Approx. 10-12 a Rivers a fi plantation So surroundings Zelenchuk River Neighbouring areas 5-12 km All the animals came back Zelenchuk observatory uence of the Psyzh and fl Linevaya Balka and Ginseng The wisents were observed to con enter the left side of Bolshoy The wisents were observed in the Tserkovnaya Polyana area near the Tserkovnaya shade-tolerant temperate coniferous forest in the Kazachya Polyana area undetermined undetermined Age and gender Age and gender - N/a 11 Adult bull Adult bull Zelenchuk observatory area All the animals came back 41 Adult bulls Adult bull Not Not side of two reserve reserve species Groups Groups to three observed observed along the consisting outside the outside the reserve left 199920002001 N/a N/a 2002 2003 1997 1998 1993 late January Chapter 4. Habitat Structure and Migration Dynamics 75

Figure 29. A Young Bull on the Forest Edge. Photo by U. Semenov. 76 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 30. The Topographic Map of the Kizgych Forestry — the Arkhyz Area of the Teberda Nature Reserve (Scale 1: 100,000). Chapter 4. Habitat Structure and Migration Dynamics 77 Many studies of contemporary scientists have been dedicated to the migra- tions of wisents in the mountains. In general, they are related to seasonal migra- tions of the animals within certain territories and associated with the depth of the snow cover in winter. During other seasons, such migrations are caused by the forced search for more comfortable conditions and the endeavors of the animals to use the localized areas in full. In the past, the native Caucasian wisents occu- pied considerable areas and could leave for valleys to spend winter there (Nord- mann, 1838) down to the present-day borders of Khamyshki, Dakhovskaya, Sto- rozhevaya, Psebay and Pregradnaya villages, as well as to the southern slope of the Greater Caucasus Mountain Range to the Black Sea shore (Latsarius, 1879; Radde, 1899; Dinnik, 1910; Bashkirov, 1939). Now, their migrations are limited to the nature conservation territories and the adjoining areas. Chapter 5. Biotopical Distribution

The complex and varied high-mountain relief of the Kizgych Valley limits the op- portunities of the wisents to undertake considerable seasonal migrations because the chine is surrounded with rocky mountain ranges on three sides, and the lower wide part of the valley ends in Arkhyz village, which also prevents active migrations of the animals. When the snow cover reaches 25–30 cm in the valley the wisents still have no real diffi culties with fi nding food but it is already almost impossible to cross the mountain passes, so the animals have to look for comfortable areas in thick conifer- ous woods within the chine. Nevertheless, the animals migrated for some signifi cant distances from time to time leaving the reserve territory. The general geographical range of the wisents in the Kizgych Chine is be- tween 1,450 and 2,800 m above sea level, but the most comfortable conditions where they are most frequently observed are between 1,450 and 1,900 m above sea level. When the weather conditions change, and there is much snow, the animals leave for fi r woods in the mouth of the Kizgych River, in the middle streams or to the confl uence of the Kizgych and the Bolshoy Zelenchuk Rivers. Solitary traces can only be infrequently found in the headstreams of the river during snowy years. During the spring and summer period, the majority of the animals move to the subalpine meadows at the upper border of the forest. They are seen at the altitude of over 2,500 m above sea level sometimes. The distribution of tree and shrub vegetation within the chine plays a consider- able role in the wisents’ choice of the feed and rest places, especially whether such wood species as the wych-elm, Caucasian maple, bird cherry tree, goat willow, ori- ental beech and others grow there. The main stations of the wisents during the sum- mer period are areas with well-developed grass cover especially from late spring to the fi rst half of summer when the plants they prefer (gramineous and legumes) are most lush and nutritious. At that time, they are most frequently observed in the avalanche cones covered with fresh greenery and fl oodplain terraces with spots of shrub vegetation. In late summer and autumn, the coarse and less nutritious grass in the forest areas gradually becomes less important in the wisents’ diet. The animals were gradually exploring the territory, developing specifi c habit patterns and getting adapted to the new life conditions in the mountains starting from the moment of their release and during the fi rst three years after that. In 1971, which was three years after the release, the wisents started walking out on Chapter 5. Biotopical Distribution 79

Figure 31. A Group of Three Bulls in the Avalanche Cone. Photo by U. Semenov. the avalanche cones where they had not been seen before. When the snow cover reached the highest level from January to March, they concentrated in fi r woods and sometimes walked out of there to the river fl oodplain and glades although they had not been to those places before. The wisents had explored the whole territory of the chine except the rocky part and sometimes crossed its borders by 1972. For example, one bull left for the headstreams of the Rapachay River that is located 8 km northeast from the reserve territory in early autumn. That wisent kept within the lake Chabakly-kel area for over a month but came back to the herd as it got cold, and snow fell. Lots of snow fell in the winter of 1971/1972, and the whole herd kept mainly on the river fl oodplain near Mertvoe Lake and on the Lebzor and Bolshaya Che- gordala glades. The wisents covered not more than 4 or 5 km a day and some- times stayed within one small area for several days during that period. In spring, starting from mid-April, the wisents began to go farther away to the upper part of the chine. Sometimes they went over to the northern slopes of the neighbouring Arkhyz Chine where they kept to the fi r woods in the pass between the Kizgych and Arkhyz Chines. The wisent habitat is wide during the summer- autumn period. The animals often went to the upper part of the forest walking out on the subalpine meadows from time to time. The daily distance covered by the animals during the warm season amounted to 4–6 km on the average. 80 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 32. A Wisent’s Lair in the Fir Woods. Photo by U. Semenov.

Figure 33. A Group of Wisents on the Sofiyskoe Sedlo Mountain Range Spurs. Photo by U. Semenov. Chapter 5. Biotopical Distribution 81 The wisents continued exploring the reserve valley and the adjoining areas but no longer gathered into one herd during the following several years. As winter came, they concentrated in the river fl oodplain except for two bulls that were going to the Psyzh isolated terrain area from time to time. Like in the previous years, the animals kept to the subalpine and alpine meadows in the Kizgych headstreams up to the Bugoy-chat isolated terrain and the Sofi yskoe Sedlo Pass in the spring-summer period until a stable snow cover settled. Small groups of bulls ascended to the Cheget-chat Mountain Range spurs above the Krest glade and Tokmak-kaya up to 1,700–1,900 m above sea level. The wisents used almost the entire territory of the Teberda Reserve Kizgych Forestry and the adjoining areas of the Zelenchukskiy forest farm with the total area of 14,000 ha in 1979. Around 15–20% of the wisent population walked out of the reserve and stayed in the adjoining territories in the Psyzh, Sofi a, Arkhyz and other isolated terrains for three to six months (sometimes up to one year and longer) during all the seasons. The other part of the animals was concentrated in the upper Kizgych Valley (blocks 55–83) and sometimes went over to the neigh- bouring Sofi a isolated terrain in small groups through the Sofi yskoe Sedlo Pass, stayed there for fi ve to eight days on the average and came back. The wisents were most frequently seen in blocks 73, 76–79, 82–83. That year, the places of their regular location during the day rest time were discovered. The

Figure 34. A Mixed Group of Wisents on the Artificial Salt Lick. Photo by U. Semenov. 82 U.A. Semenov The Wisents of Karachay-Cherkessia fi rst place was located above the Baga-tala glade at the altitude of 1,800 m above sea level on the 30-degree steep slope of the northeastern exposition in block 73 in the beech and fi r forest mixed with wych-elms and Caucasian maples with the multigrass-fern cover where valeriana alliariifolia, European goldenrod, white bog rosemary, mountain fescue and ribes prevailed in the underwood cover. The total area occupied by the wisents’ lairs amounted to around 600 square meters. That place had a good view on the surroundings, and there were rub trees – tilted fi rs; the lairs were arranged under the fi rs and were well protected from rain and snow. The wisents left signal marks, both olfactory and combined, through- out the entire territory and along its access ways. The second station of the animals was discovered approximately 900 me- ters from the fi rst one in block 71 at the upper border of the fi r forest below the wisents’ favorite subalpine grazing land to the north of the Sofi yskoe Sedlo Pass. The slope exposition is eastern, around 20-degree steep, 1,950 m above sea level. Biotope: fi r forest mixed with beeches, the average diameter of fi rs at the chest level is around 40 cm, height up to 40 m, multigrass cover with Euro- pean goldenrod and white bog rosemary prevailing. The place was followed by a subalpine tallgrass meadow. A “tochek” (stamped well-puddled section of the meadow occupying around 9 square meters, made by bulls during the mating season) was discovered there. The animals’ lairs were spread around the entire station and were located both one by one and at a distance from each other (up to 30 m). The same as on the fi rst station, they were located under the thick fi r

Figure 35. A Place of the Wisents’ Daytime Rest on the River Floodplain. Photo by U. Semenov. Chapter 5. Biotopical Distribution 83 heads, and there were various olfactory and combined marks within the territory and along the access ways. Later, in September 1980, the third station of the wisents with the area of over 500 square meters was discovered on the left side of the Kizgych River in front of the Byky rocks at the altitude of 1,510 m above sea level on the eastern exposition 10-degree steep slope in the beech and fi r forest. The height of some trees reached 40 m there. The animals’ station was located around 100 m away from the river and 30 m from the big glade. The hierarchy of lairs was not clearly defi ned here because there were plenty of places well protected from rain and snow. Although such places can be used by the animals for quite a long time, they are still not permanent. The animals change them in one to three months and move to new places for various reasons, including if a man shows up there. Besides, temporary stations where the wisents stayed for a few days could be frequently found on the sand isles in the Kizgych River fl oodplain during the summer period. That usually happens during seasonal vertical migrations. “The Caucasian wisent uses various places to stay during different seasons. According to hunters and based on my personal observations, they really like staying at the upper border of the forest in summer where the thick young birch woods, rhododendron (Rhododendron ponticum and Rh.cancasicum) and Cau- casian maple (Acer Trautvetteri) grow. Such places attract them by coolness, fresh and lush food, as well as by the lack of insects — mosquitos, gadfl ies, etc. In the morning and evening, the wisents sometimes leave for the alpine mead- ows to graze. When it rains, they go down to the spruce forest where they can fi nd a reliable shelter even from the strongest rain under the big spruces and fi rs with thick down-hanging branches and can have a rest on the dry ground covered with a layer of fallen fi r needles. I often saw one or two huge liars of wisents un- der such fi r trees growing both at the edge and in the middle of the forest, while I found even eight such lairs under a small cluster of fi rs in the Shishi Valley. In autumn wisents often stay for a rest on the mellow ground in the roots of fi rs, beeches, and other trees pulled out by the storm. A wisent turns the earth in the place where it has a rest one day after another into small dust. Sometimes several wisents choose the same place for rest so that the earth there becomes mellowed, and the grass completely stamped on the square of around fi ve sazhens”, N.Ya. Dinnik wrote (1910, p. 150) after having found the animals’ stations. During soft winters, for example, in 1992/1993, the wisents were spread al- most throughout the entire territory of the chine, and were seen both down the river and in its headstreams. Most of the time, the animals kept on the fl oodplain terraces and in small isolated terrains with smooth slopes, as well as on the ad- joining northwestern and southeastern spurs covered with fi r and spruce forests. That year was distinguished by the fact that part of the wisents spent the winter on the southwestern slopes of the Cheget-chat Mountain Range, approximately 3 84 U.A. Semenov The Wisents of Karachay-Cherkessia to 4 km above the confl uence of the Kizgych and the Psyzh Rivers. They stayed in the shade of the fi r, and fi r and spruce forests where the snow cover is always thinner than in other areas, and grazed on lichen that has an important role in the wintertime diet of wisents. The same year the wisents started to use regularly one more territory in the Baga-tala isolated terrain area where they kept till the middle of April. A small mixed herd of 9 to 11 wisents dwelled in the shade-tolerant temperate coniferous forest with single deciduous trees in the lower part of the slopes that they were leaving for the glade with uniform hard snow cover where hay stacks were put for them, and for the avalanche channel. The same as in other places, those sta- tions were rich in lichens and woody forage from deciduous trees. The animal groups seen in the midstream of the river kept within the wide fl oodplain terraces choosing areas with thin snow cover. As spring set in, and the snow started melting actively, the wisents began to move downstream to the Kizgych River mouth descending to the fl oodplain, walking from one side of the river to the other and often grazing on bark and dead grass in the avalanche cones. During that period, the snow in the lower part of the chine was melting quicker, and the animals concentrated in the Perviy Post and Krest isolated terrain areas actively grazing on the early grass plantlings. Their migration to the lower warmer areas was probably also related to the birth of youngsters when the females separated from their groups and gave birth to their offsprings in the thick shrub vegetation. Wisents could be seen at the maximum altitudes from late June to mid-au- tumn. During that period, the majority of the animals slowly moved back to the headstreams of the river where they walked from big glades — spurs of the Sofi yskiy Mountain Range — to the upper border of the forest onto the subal- pine and alpine meadows. As a rule, they stayed there till the snowfall and then they left and spread along the entire chine till the following spring. However, the autumn of 1993 was almost snowless, and a number of mixed groups of the animals continued grazing there till the end of November and were observed on the alpine meadows at the altitude of 2,400–2,700 m above sea level. As the ob- servers noticed, the Kizgych wisents had never ascended so high before. In summer of 1994, the main locations of the animals were near the upper border of the forest and the subalpine meadows. The coniferous woods there are considerably mixed with deciduous trees growing in small clusters in the middle and in the lower part of the slopes. The wisents were most frequently seen in birch woods and mixed deciduous sections, including maples, bird cherry trees, oriental beeches, etc. During that season, they were not so often seen on the river fl oodplain where they had mainly dwelled in the previous years. They dwelled there for a short term. In late November, the majority of the animals started mov- ing to the territories down the river. The extensive use of the high-mountain part Chapter 5. Biotopical Distribution 85

Figure 36. Mineral Water Sources in the Kizgych Valley Are Used by the Wisents as Natural Salt Springs. Photo by U. Semenov. 86 U.A. Semenov The Wisents of Karachay-Cherkessia of the chine by the animals was noteworthy as a peculiarity of their distribution throughout the territory in 1994. Solitary animals and small groups consisting of two to four species started showing up in the areas above the forest on the subal- pine and alpine meadows where they had not been seen before. The chine explo- ration was continued, and the animals were observed almost in all the available areas. Several mixed groups of the animals spent the entire snowless period on the mountain slopes almost not descending to the river fl oodplain. A small herd of 12 species of different gender and age dwelled in the Byky isolated terrain at around 2,000 m above sea level from mid-June to mid-September. The favorable conditions for the long dwelling of the animals in that place were created by the complex combination of the fi r woods mixed with deciduous trees, the avalanche channels covered with grass and shrubs with thick raspberry and blueberry sec- tions, as well as the forest glades and creeks nearby (Table 7).

Table 7 Seasonal Location of the Wisents in their Main Dwellings in 1994

Number of Wisents Seen Dwelling Seasons abs % Winter 24 64.8 Spring 20 54.0 Deciduous forest Summer 36 50.7 Autumn 9 36.0 Winter 13 35.1 Spring 17 45.9 Shade-tolerant coniferous and deciduous forest Summer 14 19.7 Autumn 12 48.0 Winter - - Spring - - Pine forest and pine and deciduous forest Summer 3 4.2 Autumn 4 16.0 Winter - - Spring - - Mountain meadows Summer 18 25.3 Autumn - -

Note: According to the Teberda Reserve Nature Records. Chapter 5. Biotopical Distribution 87 The conditions in the Baga-tala isolated terrain where a herd of 26 wisents was dwelling were similar. The animals were registered in the alpine and subalpine region from late July till late August. In 1995, the observers noticed that the east- ern border of the wisent habitat shifted to the Uzhum and Morkh-syrty Mountain Ranges. No migration of the wisents from the forest area of the slopes down the river, typical for the autumn period, was observed in that season of 1994. The wisents moved to the lower territories as late as in November when the snow fell. The analysis of the animal’s distribution throughout the territory of the Kiz- gych Chine and the adjoining areas where the wisents were seen from time to time as they crossed the reserve borders, showed that the animals had certain preferences depending on the types of grasses and trees, complexity and steep- ness of the relief, depth and condition of the snow cover. The animals were almost never seen on over 40–45°-steep slopes, big screes, rocky areas and creek sides with wind-broken woods and big rocks. They were seen in the pine forests a few times only. As the snow fell, the wisents migrated to the fi r woods where the snow depth did not exceed 50–60 cm even during snowy winters. The same as in the previous year, the majority of the animals kept to the slopes during the summer-autumn period and almost never descended to the fl oodplain

Figure 37. The Wisents Start Visiting Mineral Water Sources as Early as in the Beginning of Spring Some Years. Photo by U. Semenov. 88 U.A. Semenov The Wisents of Karachay-Cherkessia till the middle of autumn. Groups of the animals were most frequently seen in the Byky and Baga-tala isolated terrains. During the winter time, they concentrated in the fi r woods in the areas of Mertvoe Lake and the swamp near the confl uence of the Kizgych and the Psyzh Rivers. In spring the animals were much more often seen in the avalanche cones. A similar spread of the animals was also ob- served by the inspectors outside the reserve in the Linevaya Balka area where a group of four bulls stayed for a long time in the broad-leaved woods with a thick layer of understory trees, including aspens, maples, mountain ashes, etc. From 1996 to 1999, the territorial spread of the animals within the chine was the same. The animals mainly kept to the shade-tolerant coniferous forests and sometimes to the mixed shade-tolerant coniferous and deciduous forests, and were clearly attracted by the avalanche cones and channels, fl oodplain terraces and for- est glades rich in meadow vegetation. In winter they were more often seen on the southwestern slopes heated by the sun where the snow cover was thinner than in the rest of the chine. During the summer period and in the beginning of autumn, the animals kept to the forest glades on the slopes and mountain range spurs. Some- times they ascended to the subalpine meadows at the upper part of the forest where they were grazing for a long time amidst the crooked birch woods and at the edges of the forest covered with high grasses. In spring, more often in the second half of March and the fi rst half of April, the wisents spent the majority of their time on the avalanche cones and entered the nearby shade-tolerant coniferous woods only when it was windy, rainy or snowy. Later, in the end of spring and the beginning of summer, they went down and grazed in the valley or on the fl oodplain terrace glades as the grass started to grow. The spread of the wisents within the territory of the Kizgych Mountain Valley was mainly linked to the Tokmak-kaya, Byky, Baga- tala and Bugoy-chat isolated terrains. So the wisents developed a certain system of using the high-mountain land- scapes of Arkhyz as a result of their dwelling in the mountain ecosystems for over 20 years. Today their seasonal and daily migrations help them to use almost all the varied biotopes of the Kizgych Chine except for its rocky part. The snow cover distribution has become the main reason for the seasonal migrations, while the daily movements are caused by the search for more comfortable places for rest and grazing, as a rule. In such conditions, the migrations do not generally exceed a few kilometers and sometimes can be even reduced down to a few hundreds of meters. In the recent decade, they have almost no permanent dwellings. They consistently migrate from one place to another not covering any huge distances but shifting through a large number of stations. As a result, the spread of the animals along the biotopes within a year is of clearly seasonal nature as follows: Alpine (including subalpine) meadows — the wisents start showing up there as early as in May following the snow melting. That is a typical summer grazing place for the animals. In hot weather they graze there during the early morning hours and Chapter 5. Biotopical Distribution 89 leave the open spaces by 9–10 a.m. hiding from the summer heat under the shade of the coniferous woods, then they walk out again after 7–8 p.m. and graze till dusk slowly moving in the direction of the forest. Around 8% of cases when the wisents were seen by people happened in the alpine and subalpine areas (Fig. 38). The mixed coniferous forests play an extremely important role in the life of the Arkhyz population of wisents and are used by them throughout the year. During the summer period, the animals fi nd coolness and shade there and spend almost all their time of rest in such places. In winter the mixed coniferous forests are the center of the biotopic spread of the wisents within the chine territory. During the fi rst years after the release of the animals, they fed on hay and the availability of natural food was not of critical importance for them, while in the follow- ing years, the snow cover distribution and thickness became the drivers of their choice of places for grazing and rest. The snow level is considerably lower in the coniferous forests, especially in the fi r and spruce woods where part of the snow is settled on the tree branches, than in the open spaces. Wisents fi nd food and shelter from snowstorms and cold winds under the thick heads of the mixed coniferous woods (around 9% of cases when the wisents were seen). Even mi- nor distances covered by the animals when they are looking for food are routed through the coniferous woods because the animals can always fi nd a way there through areas with thinner snow cover. Pure birch woods, as well as beech woods, pine forest areas, and spruce woods are rarely visited by the wisents. Birch woods do not occupy any large spaces in the Kizgych Chine, growing mainly on the fl oodplain terraces alongside the river and forming spots of various shape and area that usually do not exceed 0.5–0.9 ha. The snow starts melting there early in spring, and the thaw holes become covered with the fi rst grass that the wisents eagerly graze on. As a rule, the wisents do not stay in such areas for a long time grazing and having a rest there for a short term. During the winter period, quite a lot of snow accumulates in such biotopes and reaches 1–1.5 m height by spring. The wisents try to avoid such areas, crossing them if they have no other choice only. We have never seen the wisents having sta- tions there in winter for the entire period of observing the animals. Around 0.8% of the cases when the wisents were seen only happened in the birch woods. The same is true for pure beech woods (0.4% of cases), pine woods (0.4% of cases), and spruce woods (0.2% of cases). Although the wisents eagerly graze on their young shoots, such biotopes are just intermediate locations on their way to the places of grazing and rest, which is probably caused by their poor protection characteristics in the fi rst place. In winter they are inconvenient, cannot protect them from cold wind or rain and are easily seen through. In summer the wisents keep at higher altitudes in the subzone of the subalpine and alpine meadows avoiding gnats. The animals are more frequently seen in aspen woods, around 3% of all cases. The aspen is one of the favorite food types for wisents; they eagerly graze on its young 90 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 38. Biotopical Spread of the Wisents in the Teberda Reserve Kizgych Forestry (Based on the Observations from 2000 to 2010).

shoots, leaves and bark regularly showing up in the aspen forest sections — 3% of all cases. Fir woods hold a special position among the mono-dominant forests with the 3% share in the total cases when the wisents were seen. One can see the animals there the whole year round. The relatively low percentage is due to the small areas occupied by the pure fi r woods. The mixed deciduous forest is visited by the wisents much more frequently (4% of all cases) than pure mono-species woods. It occupies the Kizgych River fl oodplain terraces and runs along the entire chine wrapping the glades in the widest areas on the both sides of the river. The wide variety of species on rela- tively small areas increases the chances of the wisents to fi nd food in such sta- tions. Lairs of the animals can be found in mixed deciduous forests unlike in beech, birch or pine woods. However, our observations showed that the wisents tended to leave such biotopes trying to move to the fi r woods or to the areas where the forest was mixed with coniferous species. The wind does not pass well through mixed deciduous forests located at relatively low altitudes, so the concentration of gnats, mosquitoes and gadfl ies is very high there during the spring-summer period. That is why the wisents coming there in summer do not stay in such places for a long time. In winter the snow cover there is thicker than in coniferous woods, so the protective properties of such forest drop, which also drives the animals away to other territories. The mixed coniferous and deciduous forest is one of the most comfortable biotopes for the wisents in the Kizgych Chine — over 11% of the cases when wi- Chapter 5. Biotopical Distribution 91

Figure 39. As the Wisents Walk the Same Ways All Over Again They Eventually Stamp Wide Paths that Are Then Used by Other Animals. Photo by U. Semenov. 92 U.A. Semenov The Wisents of Karachay-Cherkessia sents were seen. They occupy large areas and thus allow the animals to stay not far from the forest glades and to fi nd food and shelter from bad weather without taking up any long-distance migrations. However, it should be mentioned that the animals prefer such types of coniferous forests that have a considerable share of Nordmann fi rs. Those trees may reach over 70–80 cm diameter and 35–40 m height. Spots of fallen needles are accumulated under their heads where it is dry during the rain, and the snow cover is thick even in especially snowy years. Other species in such forests allow the animals to fi nd food without consider- able movements and make lairs in the same place. The wisents use such stations throughout the year, and some of them even one year after another. The mixed coniferous and deciduous forests occupy the area alongside the lower part of the northern and southern exposition slopes and lie at a considerable distance from the cold winds blowing at the bottom of the chine. They can always shelter the wisents from them in winter. At the same time, there are a lot of areas easily accessed by the wind where the wisents can hide from gnats and heat in sum- mer. The mixed woods commonly climb 2/3 of the slope height in the shape of a tongue which makes them more attractive for use throughout the entire year. Roads are used by the wisent infrequently, less than 1% of the cases. For the fi rst years after the release of the animals, the vehicles carrying hay left a wide track in the deep snow. The wisents frequently used some of its sections for walking. After the feeding of the wisents was stopped, no frequent trips were made to the Kizgych except for carrying materials for security station repairs, hay for horses, salt for artifi cial salt licks, etc. The animals often crossed the track made by a vehicle moving in the deep snow as they migrated from one area to another but in some cases the wisents walked in it for up to 200 m probably looking for a better place to cross it and move on. Then they left the road and went their way. In fact, the wisents did not walk out on the road on purpose but if they came across it they used its small sections to make their movement easier. Meadows and forest glades are similar stations in many aspects but they dif- fer in terms of area and location. The fi rst category includes relatively big areas (over 1 ha) located in the wide river fl oodplains. The second category includes small territories fully or partially surrounded by the forest and located on the fl oodplain terraces and mountain slopes. Both the fi rst and the second types of stations are used by the wisents throughout the year almost equally: around 10% and 9% of the cases, respectively. The observations of the animals showed that the wisents still have certain pref- erences depending on the season. For example, the wisents show up more fre- quently on meadows starting from October and almost till summer. The autumn migrations are apparently related to the reduced number of gnats on the fl oodplain and at the riverside as a result of the temperature lowering. The animals stay there for the entire winter but no grazing traces are found where the snow depth exceeds Chapter 5. Biotopical Distribution 93 40-50 cm. At that time, the wisents switch to the woody forage, and the meadows are not used by them as grazing stations any longer. Forest glades, especially those located above the fl oodplain terraces are attractive for the animals as comfortable places during the summer period. It is always cooler, and the blood-sucking insects are not so abundant there. They continue using such places until deep snow falls. When winter sets in, the wisents walk out on such slopes less frequently and stop going there at all as the snow cover becomes thicker. Gullies and draws are used by the wisents very infrequently, mainly as a way to other areas (around 1% of cases). It should be mentioned that such stations are inconvenient to wisents for long-term stays in general. For the entire period of observations, we have never seen that the animals stayed there for any long time. In one case, the wisents were attracted by the crab apples rolling down the slopes of a small gully, in other cases – by the traces of the animals that had walked there. Paths are an important and integral element of the animals’ life in the moun- tain conditions. The complexity of the highly broken ground sometimes restricts movements to specifi c paths, and the animals have to use the same routes for their local walks, long-distance runs, seasonal and other migrations. For exam- ple, the wisents used the pass to the Sofi a isolated terrain that lay along the slope of Sofi yskoe Sedlo Mountain and two paths to access the mineral water source for many years, the Nordmann fi r that the animals regularly used for rubbing had only one access path, etc. Notwithstanding the small percentage of cases when the wisents were seen on the paths (total around 2%), they are extremely important for the animals’ life and play a key role in the local movements of the wisents driven by their feeding behavior or by the search for better places to stay. Avalanche cones are most frequently visited by the wisents from February till late March-early April although they can be seen there all year long — 16% of the cases1. The animals use mainly the lower part of the avalanche channel where the maximum volume of highly compacted snow is accumulated, and young de- ciduous survivor trees of 2–4 m (birch, maple, mountain ash, bird cherry trees, etc.) grow on the knaps. The upper part of the avalanche cones — their origin places and channels lying in the rocky mountain combes — is not used by the animals. Notwithstanding the fact that the wisents frequently use the avalanche cones, they still should be considered an important intermediate grazing sta- tion but not a main one although their importance in the animals’ diet abruptly increases in some years. The observations showed that the wisents usually use them when walking to the grazing territories in a temporary location area but do not go to them on purpose covering any considerable distances. The nature of

1 The high percentage of cases when the wisents were seen in that biotope is partially caused by the observation peculiarities in the mountain conditions when the wisents can be more often and easier seen against the light background of the avalanche at a long distance, which can cre- ate a wrong perception of their preferred stations — author’s note. 94 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 40. The Wisents’ Rub Tree Used by Them for a Number of Years. Photo by U. Semenov. Chapter 5. Biotopical Distribution 95 the avalanche channel used by the wisents is different — it depends mainly on the snow volume. In snowy winters, the number of avalanches and the size of the cones go up abruptly, which causes considerable destruction of the existing plant associations and drives the renovation and rejuvenation of the tree and shrub vegetation structure. During the fi rst one or two years after such winters, such biotopes are infrequently visited by the wisents as they lose their grazing attractiveness because the old vegetation is destroyed, and the new vegetation is growing very slowly, while such areas are used by the wisents more actively further on. Leaves and young shoots of the thick birch, maple, raspberry and other vegetation that appear in the avalanche cones every year are an additional reserve of woody forage making such places more attractive for the animals and are actively eaten by them not only in early spring but also during the spring- summer period. As a result, 27–48% of the trees growing there (maple, birch, mountain ash, etc.) had their bark and shoots damaged. The biotopes close to water such as riversides, lakeshores and small swamps are used by the wisents differently (11, 1 and under 1%, respectively). The ani- mals came up to the lake and the swamp mainly when the meadow grass coars- ened and burnt in late summer and autumn while the grass on the shore of the drying lake located in the lower part of the chine and on the tiny swamps along the small creeks remained green and lusher. The riversides and areas along big- ger creeks were used by the animals almost all the year round. Mixed deciduous forests consisting of such species as aspen, alder, willow, mountain ash, bird cherry tree, etc. form favourable grazing stations for the wisents and are actively visited by them almost all the year long. The various species composition of the tree and shrub vegetation together with small glades covered with thick and lush grasses make the riversides one of the most attractive grazing biotopes for the animals. Their role is especially important during the autumn and spring periods. Mineral water sources and artifi cial salt licks are the places where the wisents satisfy their requirement in minerals. They are an integral element of their daily migrations and have been used by them almost equally during the recent decade — around 5% of the cases both. The animals’ movements to mineral sources are most active from early spring till late autumn. It should be noted that the animals access the salt licks during the winter period, as well although much less frequently than during the rest of the year. Such important components of the mountain biotopes as crooked forests, high-mountain woodlands and rocky areas are almost not used by the wisents. For the entire period of our observations, we have never seen the animals in such stations. They also avoid crossing areas with large rock debris and wind-broken woods in the mudslide or avalanche areas. Chapter 6. Diurnal Activity

“The Caucasian wisents spend a considerable part of the day sleeping or doz- ing in their stable or just somewhere in the forest and then go grazing or drink- ing water about three-four hours before the sunset. At that time, they often walk out on the glades or forest edges especially preferring birch woods at the upper border of the forests. In the morning, when the sun is more or less high in the sky, they go back to the forest. However, the wisents sometimes graze in the afternoon, too either walking out on the glades or mountain meadows or stay- ing within the forest”, Dinnik wrote describing the diurnal activity of the native Caucasian wisents (1910, p. 151).

Figure 41. A Wisent on the Salt Lick in the Early Morning. Photo by U. Semenov. Chapter 6. Diurnal Activity 97

Figure 42. Diurnal Activity of the Wisents in Different Seasons.

The diurnal activity of the wisents reacclimatized in Arkhyz is mainly de- termined by their feeding behavior and differs based on the season depending mainly on availability of food within the period (Fig. 42). As it was noticed as

Figure 43. When It Is Cloudy, and the Sun Is not so Active, the Wisents Often Have a Rest on the Meadows and Slopes not Far from Their Grazing Places. Photo by U. Semenov. 98 U.A. Semenov The Wisents of Karachay-Cherkessia early as in the beginning of the 20th century (Dinnik, 1910), the wisents’ feed- ing periods are broken with rest intervals during the day. In summer when the animals have enough food, their feed and rest phases are distinguished more clearly. At dawn the wisents start grazing actively (33% of the cases when the wisents were seen) for several hours but the majority of them go for a rest at 10–11 a.m. already leaving open spaces for the wood shade. The time when the wisents leave the meadows and glades can shift back or forth depending on the weather. On hot days when the ambient temperature reaches 20–25 degrees by

Figure 44. The Wisents Visit Salt Licks Even at Night during the Spring-Summer Period. Photo by U. Semenov with hunting camera.

10 a.m., the wisents leave open spaces and continue grazing in the forest where the temperature is signifi cantly lower. The next most active phase starts at 5–6 p.m. and continues till the twilight. We had no opportunity to observe the wisents grazing at night till 2012. So our data (6% of the cases) were collected during the late twilight (around 9 p.m.) when the animals could still be observed visually. It should be noted that the wi- sents sometimes graze at night during the winter period. Although it is diffi cult to visually observe the grazing animals in such conditions, the paths of traces on the snow discovered in the morning helped to fi nd the newly browsed tree bark and branches. Hunting cameras help to collect more accurate data. Chapter 6. Diurnal Activity 99 The wisents may be at the stage of “passive” feeding between the morning and evening activities in the forest or have a rest under the heads of thick fi r woods. The nature of such feeding is as follows: the animals walking out to open spaces of the subalpine and alpine areas or forest glades and meadows disperse and move slowly in one direction cropping grass as they walk. Sometimes the wisents grazing on small glades stay there for a longer period and then go to another place. They stay longer in localized areas of the forest zone because they graze on woody forage and bark there which takes more time. As you can see in the graph (Fig. 42), the diurnal activity of the wisents during the autumn-winter period differs signifi cantly from their spring-summer feeding behavior. The number of activity phases of the animals reaches fi ve or six per day. The wisents keep grazing for almost all the daylight hours (29% of the cases in the morning, 38% in the afternoon, 33% in the evening) pausing for a rest for quite a time. The higher the snow cover is, the more time the animals need to satisfy their hunger. The snow cover of 15–20 cm has almost no relevant impact on the feeding behavior of the animals, but when it reaches 60–80 cm their mi- grations are considerably limited, and they have almost no opportunity to extract grass from under the snow. The majority of their food during such periods is the wood forage and bark of trees and shrubs. Abiotic environment factors infl uence the diurnal activity of wisents consider- ably. Although the animals were seen grazing almost in any weather the majority of cases was registered when the sky was cloudless — 42% of the cases (Fig. 45). On cloudy days, the animals’ activity was decreased somehow but still re- mained at a quite high level of 29–30%.

Figure 45. The Wisents’ Activity in Different Weather. 100 U.A. Semenov The Wisents of Karachay-Cherkessia

46 50

47 51

48 52

49 53 Figures 46–53. A Group of Four Species Visited the Mineral Water Source and Spent Almost the Whole Day there from 9:27 a.m. (Fig. 46) till 8:20 p.m. (Fig. 53). Photo by U. Semenov with hunting camera. Chapter 6. Diurnal Activity 101 When it was raining or snowing, the wisents’ grazing activity was signifi - cantly lower although the highly variable weather in the mountains causes the animals to adapt in certain ways while its abrupt change frequently leads to al- teration of the biotopes. During the cold season, the animals leave open spaces, while in summer they almost never stop grazing even in case of drizzling rain. In autumn and winter, the animals try to leave for the coniferous forest in advance, especially female wisents with calves of the current year. The majority of the animals registered during such period formed small groups (up to three species) consisting mainly of bulls (around 19%). The wi- sents showed the lowest activity when it was snowing — around 11% of all the cases. However, based on our observations, the adult animals continued graz- ing even in open spaces as it started slightly snowing. The groups that included calves of the current year left open spaces for the coniferous forest and moved almost without stopping. The paths of the wisents’ traces in the woods showed that the animals that have left open spaces sometimes continue grazing in the forest covering short distances within localized areas of the coniferous forests. Chapter 7. Diet

The diet of the wisents during their fi rst year of reacclimatization in Arkhyz was not really varied. During all the seasons of that year, the wisents grazed on bark and small branches of the goat willow, mountain ash, aspen, alder, bird cherry tree and hazel, as well as the Caucasian whortleberry and rhododendron, and various grasses with gramineous species prevailing. In the late 19th – early 20th centuries, scientists studying the diet of the native Caucasian wisent discov- ered around 40 plants that are most preferred by the animals throughout the year (Dinnik, 1884, 1890a, b, 1910; Filatov, 1910, 1912; Bashkirov, 1939). The variety of species in the diet of the wisents in the Kizgych Valley resulted from the location of the animals during specifi c periods. A part of the brought in animals that stayed in the subalpine territories in the beginning of winter grazed on various grasses of the subalpine and alpine meadows. But the food variety was not so wide there because the domestic cattle had been grazing there till the early 80s, so the coloured fescue dominated in the diet of those wisents. The lack of snow cover allowed the wisents to regularly go to the forest where they ate rhododendron, fi r needles and small branches of deciduous trees. Although the winter was not snowy, such type of feeding turned out to be insuffi cient for the animals of that group, and they looked exhausted and weak in spring. Another part of the animals that spent that winter in the valley was actively eating hay, plus the composition of the woody forage in their diet was more var- ied due to the wider range of tree and shrub species growing in the lower part of chines. As for grasses, they grazed on the reed grass, horsetail and various dead grasses that they could easily fi nd under the trees. In spring snow melted fi rst on the fl oodplain terraces, so the wisents of that group started grazing on fresh grass earlier, which also helped them to quickly recover after their fi rst winter. In sum- mer their diet often included the cock’s-foot and blackberry, while the woody forage prevailed during the winter period. Notwithstanding the little snow that fell in the winter of 1968/1969, some of the wisents survived it with diffi culty. The hay stocked in advance played the key role in their survival. It was doubtful whether the wisents would be able to survive on their own in more severe snowy winters. Further observations proved that supposition to be false, but still the stocked hay was the main condition of Chapter 7. Diet 103 the wisents’ survival during the fi rst winters. Ten artifi cial salt licks were created the same year. 10–25 kg of salt bricks were put into each salt lick. The animals spent the following winter in the lower part of the Kizgych Val- ley. None of them stayed at the upper border of the forest or in the alpine zone. The wisents concentrated in the lower part of the chine and actively grazed on the dead cock’s-foot, reed grass, coloured brome and horsetail grasses that grow abundantly on the river fl oodplain. As for the tree forage, they ate branches and bark of the goat willow, alder, mountain ash, bird cherry tree and hazel. In winter they eagerly ate mountain ash berries, while from mid-December till early April, they grazed on the stocked haystacks from time to time. The most diffi cult period of that year was the beginning of spring when the tem- perature in Arkhyz dropped, and snowfalls started. The wisents could not reach the hay because of the ice-covered ground and the thick snow cover of up to 70–150 cm in some places. Their main food at that time was the bark and branches of deciduous trees. The animals lost lots of weight but the diffi cult conditions did not last long, and the wisents could graze on fresh grass shortly after. When green vegetation appeared in the wisents’ diet they started consuming salt though they had never touched it in winter. From late April till early May, the animals started showing up on the subalpine meadows where the Caucasian cephalaria, coloured fescue and other species were added to their diet. It was noticed that they ate young shoots of currant, blackberry, coloured brome, reed grass, coloured fescue, bark of deciduous trees, as well as pines, fi rs and spruces in summer. The wisents’ diet became more varied in the course of time, so the animals’ traces more frequently led to elms and bird cherry trees with nibbled bark in the winter of 1971/1972. The animals certainly preferred them when they had a choice but they continued feeding on the bark of other trees, as well. Reserve employee A. Batchaev noticed that when it was very cold the wisents more ac- tively grazed on thin shoots biting them off birches, spindles and hazels rather than on the new bark of willows, mountain ashes and aspens that is hard to tear off the trunks because of the frost. The animals chose young trees with the trunk diameter not exceeding 8–12 cm to nibble their bark except for elms as they took the bark off their trunks of 20 cm and more reaching the height of 2 meters. According to the observers’ reports, the wisents rarely ate the bark of growing pines and aspens but fallen trunks were cleared from bark totally by them. That year the wisents infrequent- ly grazed on the stocked hay. Many cases were registered when they approached the hay stacks but according to the observers, they were pulling and throwing the hay round with their horns rather than eating it. Their traces by the salt licks were fi rst found in mid-April when the ungulates started switching to fresh grass. In spring of that year, the wisents continued grazing on coarse forage and used the hay stacks more frequently that in the beginning of winter. At that time they 104 U.A. Semenov The Wisents of Karachay-Cherkessia mainly nibbled currant shoots and sometimes ate moss and lichen. They grazed on the dead grass, generally gramineous plants, from the previous year and new grasses in thaw holes. Although during the summer-autumn period the wisents feed mainly on grass, their diet includes a considerable percentage (up to 40%) of tree branches on the regular basis. In winter, especially in snowy years, the share of coarse forage in their diet increases signifi cantly. For example, even during the winter season of 1973/1974 when the conditions were favourable, the specifi c share of coarse for- age in the wisents’ diet reached 70–80%. They fed mainly on bark, new branches and shoots of birches, mountain ashes, aspens, spindles, goat willows, alders, wych-elms, bird cherry trees, pines, fi rs, hazels, and other trees and shrubs. They ate hay less frequently during that winter although they regularly approached the stacks. As for the lush food, they eagerly grazed on shoots of the blackberry, horsetail and reed grass. In the fi r, and spruce and fi r woods, they fed on the mountain fescue eating moss and lichen sometimes, mainly the beard lichen. That winter it was registered that the wisents entered a salt lick, which had not been observed in the previous winters. During the following winter, they visited the hay stacks more actively, and the bulls ate the hay more often than the females. The wisents dug snow where its

Figure 54. Traces of the Wisents Feeding on the Young Aspen Bark. Photo by U. Semenov. Chapter 7. Diet 105 height did not exceed 20–30 cm reaching blackberries, their favourite food, while they switched to branches and bark in the territories with thicker snow cover. The species composition of trees and shrubs in the animals’ diet was almost the same as during the previous years but the honeysuckle occupied the smallest share in the coarse forage they ate. Not a single case of them showing up on the salt licks was registered. They fed on the stocked hay starting the end of February. From early spring till the beginning of summer, the share of green food in the wisents’ diet grows sustainably. They rarely drink water during such periods substituting it with lush young greenery. The role of leaves and new branches of trees and shrubs grows starting from mid-June. The animals become choosier about food — they more frequently feed on coarse forage and eagerly visit salt licks. In 1974, the wisents spent the winter easier, so that they were fl eshier dur- ing that spring than during the previous one. The distinctive feature of the winter of 1975/1976 was that the wisents used the stocked hay more often than in the previous years because there was a lot of snow. The snow cover reached 1 m 40 cm in the river mouth. The animals could not move in the thick snow and had to walk in the sledge track stopping by the hay stacks for a long time. During that period, the road leading to the feeding grounds was being cleared in the lower territories of the Kizgych Chine (surroundings of the bog lake, the suspension bridge over the river and the hay cutting glade). The wisent keeper brought the hay by tractor and put it at Y-trees. Their spring diet did not differ much from that of the previous springs. In late March, the wisents almost stopped eating the hay, while the share of trees, shrubs and new grass in their diet increased considerably. The wisents spent that winter with little snow satisfactorily. The female species looked much fl eshier than the bulls in spring. They had been more actively looking for food than the bulls and started visiting hay stacks earlier. It is interesting that those were usually the females that destroyed the fencing around the stacks, too even leaping over such fencing at a height of up to 1 m 70 cm sometimes during the periods of famine. The permanent salt licks were replenished with salt as the wisents were eating it out. They were visited most frequently from May to October. In July around 36 tons of hay was stocked for winter — it was stacked and fenced in fi ve points (the insolated terrains of Perviy Post, Krest, Chegordala, Grushevaya Polyana, and the suspension bridge surroundings). There was little snow during the following winter of 1976/1977, so the wisents fed mainly on natural food and did not eat the hay. In general, their diet of that year did not differ much from that of the previous years. In summer the observ- ers noticed that the wisents visited the mineral water source in the headstreams of the river. Although they actively consumed salt on the artifi cial salt licks, too, they were seen more frequently at the mineral water source during the spring- summer period. The fact that they started using natural salt sources more after 106 U.A. Semenov The Wisents of Karachay-Cherkessia a few years spent in the Kizgych Chine Mountain Valley proved the continued adaptation of the wisents to their new environment. The animals were gradually changing the life style they had been used to in the plain forests where they had been taken from, in line with the new more diffi cult mountain environment. Bioengineering works still continued more than 10 years after the beginning of the wisents’ reacclimatization in Arkhyz when almost the entire herd was re- plenished with a new generation of wisents born in the mountain environment. In 1979, the reserve staff stocked 15 tons of hay that was totally eaten by the wisents. Their impact on the tree and shrub vegetation increased considerably, so a special check was carried out to evaluate it in the end of May. A distance of 15 km was routed along the right side of the Kizgych River from the lower part of the chine to its upper part. The width of the check territory was 10 m within which all the damages made by the wisents to trees and shrubs were counted, and then the number of counted nibbles was calculated per 10 km of the route. Two checkers carried out the check along the path. One of them examined the left part of the routed territory, and the other one — its right part (Table 8). Although seasonal differences are well distinguished in the wisents’ diet, the species composition of the food consumed by the wisents during certain periods became more or less consistent in the course of time. The spring-summer diet of

Figure 55. Traces of the Wisents Feeding on the Young Tree Bark in the Avalanche Cone. Photo by U. Semenov. Chapter 7. Diet 107 the animals almost did not differ from year to year in the early 90s because the wisents had nearly totally explored the whole variety of the plant associations growing in the Kizgych Chine and the adjoining territories. The winter diet of the animals was also similar from year to year but could still differ consider- ably depending on the snow cover depth and the ambient temperature. The role of specifi c plant species increased signifi cantly during some years though the general species composition remained almost the same. Another check of the damaged vegetation was carried out in 1980 in order to study the role of specifi c species of trees and shrubs in the wisents’ diet (Table 8).

Table 8. Registered Feedings of the Wisents (According to the Teberda Reserve Nature Records of 1979/1980)

May 21, 1979 May 21, 1980 Plant Species Per 10 km Per 10 km Total Nibbles Total Nibbles Distance Distance Bird cherry tree 53 35 73 49 Grey alder 48 32 92 61 Goat willow 46 31 64 43 Caucasian mountain ash 41 27 44 29 Aspen 26 17 29 19 Wych-elm 22 15 5 3 Birch 4 3 4 3 Norway maple and Caucasian 4364 maple Spindle - - 7 5 Nordmann fir - - 4 3 Rose - - 1 1

Note. The measuring unit for shrubs was a nibble on one shrub irrespective of the number of damaged shoots; for trees — any damage.

However, such a type of check could not objectively show the general spe- cies composition of the consumed and preferred food because the route lay 108 U.A. Semenov The Wisents of Karachay-Cherkessia along the river fl oodplain where the species ratios within the wood structure were determined by the type of their growing environment. Actually, the check studied sections of the routes of the animal groups crossing the fl ood- plain and using the food that they could fi nd there and not that they really preferred. So such checks did not cover the species that did not grow on the fl oodplain terraces within the check route or that grew there in a negligibly small quantity which resulted in a wrong perception of the preferability of the studied food.

Figure 56. Traces of the Wisents Feeding on the Young Tree Bark. Photo by U. Semenov. Chapter 7. Diet 109 The distinctive feature of the winter season of 1979/1980 was that starting from February small groups of the animals were registered on the avalanche cones, where they ate young shoots of birches, bird cherry trees, willows and mountain ashes. The wisents had never been noticed there before, so their explo- ration of new biotopes could be deemed the continued adaptation of the popula- tion to the mountain environment. That year the animals started eating hay as early as in mid-December and con- tinued using it till the end of February. At the same time, they used every chance they had to feed on the new bark of pines and on beard lichens that they found on broken or snow pressed branches. The woody forage prevailed in the wisents’ diet till mid-May and then was replaced with the growing share of grasses that became their main food during the summer-autumn period. The animals feed on coniferous trees less than on deciduous trees during any season, and wisents usually nibble bark of the fi rs and spruces from their bottom, while that of the pines — from the trunks and branches of the fallen trees. The wisents’ diet during the winter and early-spring period of 1981/1982 was similar to that of other years with the main food being branches and bark of aspens, mountain ashes, spindles, bird cherry trees, maples and hazels. As for the conif- erous species, the animals eagerly grazed on bark of young fi rs, while in March they were seen eating bark from a pine felled by the wind. Trees and shrubs in the places where the animals fed were noticed to be considerably damaged. Fourteen tons of hay was stocked and put in different locations within the river fl oodplain to feed the animals during the winter period. Solitary animals and groups were seen visiting the hay stacks from January 4 till March 30. In January the animals ate around 0.5 t of hay, in February and March — 4.5 t and 4 t, respectively. The wisents occasionally visited the scattered stacks and ate the remaining hay in early April, as well. They satisfi ed their requirement for miner- als with the salt put into the artifi cial salt licks. They were seen there throughout the year but the most frequent visits were registered from the second half of May till August. One more check of the animals’ feeding traces was carried out to continue studying the impact of the wisents on the vegetation in the Kizgych Valley and the adjoining territories (Table 9). That time the data were collected by the track- ing method following the traces of a wisent herd consisting of 19 species during one day (February 13, 1982). Such method showed a more complete picture of the wisents’ impact on the tree and shrub vegetation though it also had its draw- backs related to the few samples of checked territories and the small number of observations. The check route lay in the area of the Perviy Post isolated terrain in the territory of 20х500 m. The diameter of the damaged trees was within the range of 3 to 22 cm. 110 U.A. Semenov The Wisents of Karachay-Cherkessia Table 9. Registered Feedings of the Wisents on Trees and Shrubs in February 1982 (According to the Teberda Reserve Nature Records)

Registered Damages Species pcs % 1 Maple 93 45.4 2 Mountain ash 64 31.06 3 Fir 27 13.10 4 Aspen 11 5.34 5 Hazel 8 3.88 6 Bird cherry tree 2 0.97 7 Spruce 1 0.48 Total 206 100

Note. Any bark feeding trace on one tree irrespective of the damage extend was deemed one measuring unit.

In winter the wisents fed on branches and bark of mountain ashes, spindles, bird cherry trees, maples, aspens, and hazels. Alongside with bark and end branches, they also ate berries of mountain ashes. The wych-elm was not cov- ered by the damage counting check though the wisents used every chance to feed on bark of those trees. As for the coniferous species, the wisents’ feeding on bark and needles of young fi rs was registered. When the wisents crossed the river they stayed in the fl oodplain areas of the mixed deciduous woods with the alder prevailing among other species. They grazed on its bark from young trees and thin branches. As for the shrubs, the animals’ nibbles were registered on the rhododendron, blackberry, blueberry, currant, etc. The reed grass and horsetail prevailed among other grasses in their diet during that period. During the summer period, their grass diet consisted mainly of the cock’s-foot, galega, couch grass, Caucasian cephalaria, tall millet grass, etc. They less fre- quently ate the mountain fescue, reed grass, cat’s-tail, campion and very rarely the sorrel and horsetail. As for the shrubs, they preferred the blackberry and goat willow. They ate leaves of the maple, bark of the aspen, new shoots of the horn- beam, bark and shoots of the bird cherry tree, preferring them among other trees. As winter came, the wisents started feeding on bark and new shoots of the wych-elm, hazel, spindle, bird cherry tree, mountain ash, alder and aspen again. Several times they ate bark and needles of the young fi rs and yews. As for the wild fruit trees, their nibbles were found on bark and shoots of the cherry plum and on end branches of the young rose. The wisents eagerly ate fallen fruit of the wild pear and cherry plum in the area of the Grushevaya Polyana isolated terrain (Bobyr, 1992). After the twenty years that the wisents spent in the Kizgych Val- Chapter 7. Diet 111 ley, their impact on the plant associations of the chine became noticeable. The fi rst signs of the shrub vegetation oppression appeared, especially in relation to spindles. Wych-elms were seriously oppressed among the tree species. Many of the trees started wizening and dying because the wisents regularly fed on their bark and branches, especially in the places where they spent winters. The wisents’ diet studies showed that young species aged two to three years need up to 28.5 kg of food per day during the summer time. The animals aged over three years consume around 32 kg in the same conditions. In winter their feeding requirement is almost four times lower (Gebezynska, Krasinska, 1972). According to other data, a wisent eats 20 to 50 kg of raw food per day depend- ing on the age and condition of the animal in summer, and almost two times less during the winter season, returning up to its previous consumption level in spring, which is caused by the changing metabolism processes in the animals’ organisms (Zablotskaya, 1957; Kalugin, 1958; Kholodova, 1989; Kholodova, Belousova, 1989). The experience of S.G. Kalugin (1968), who studied the wisents’ diet in the Caucasus for a long time, showed that wisents needed at least 3.5 ha of pasture lands per one species with the glade to forest ratio of 2:1.5 for comfortable liv- ing during the spring-summer period. Unfortunately, there are not enough data

Figure 57. Wild Ungulates Eat Away the Ground of Artificial Salt Licks down to the Hard Rock. Photo by U. Semenov. 112 U.A. Semenov The Wisents of Karachay-Cherkessia to carry out detailed comparative analyses of the diets of the native Caucasian wisents and the present day lines because the former are very poorly studied. Nevertheless, as early as in the beginning of the 20th century, scientists discov- ered the frequently eaten plants (Filatov, 1910, 1912) not exceeding a few doz- ens in total. It was specifi ed that the list of the eaten food was far from full. It was discovered way back by N.Ya. Dinnik (1899, 1910) that the distinctive feature of the wisents’ diet was that the animals eagerly ate ferns, unlike other wild un- gulates and domestic cattle, even in summer when there was no lack of food and used any chance to enjoy bark of the mountain ash and elm. In the course of time, the diet of the wisents brought to Arkhyz probably did not differ much from that of the native species. The data on their feeding collect- ed during all the seasons from 1996 to 2000 (Semenov et al., 2001) showed the following composition of the most frequently eaten plants: goat willow, moun- tain ash, bird cherry tree, Norway maple, hazel, rose, birch, spindle, aspen, al- der, Nordmann fi r, tall millet grass, fescue, horsetail, cock’s-foot, fodder galega, Tatarian cephalaria, Caucasian lady’s mantle, red clover, reed grass, hogweed, Caucasian blackberry, raspberry, butterbur, Caucasian sedge, meadow vetchling, etc. (Table 10).

Figure 58. If Artificial Salt Licks are Created in the Tree Roots the Animals Will Eat Away the Bark and Soil within a Radius of up to Two Meters. Photo by U. Semenov. Chapter 7. Diet 113 Table 10. Plants Eaten by the Wisents in Arkhyz (According to the Nature Records of 1968 /1999 and Based on the Field Observations from 2000 till 2012)

Common Name Latin Name Eaten Parts Cherry plum (myrobalan 1 Prunus divaricata Ledeb. fr.l.sh.r. plum) 2 Tall millet grass Milium schmidtianum C. Koch. l.s. 3 Coloured reed Calamagrostis varia (Schrad.) Host l. 4 Reed grass C. arundinacea (L.) Roth l. 5 Cock’s-foot Dactylis glomerata L. l.s. 6 Field brome Bromus arvensis L. l.s. 7 Smooth meadow-grass Poa pratensis L. l.s. 8 Mountain fescue Mountain fescue M.B. l.s. 9 Meadow fescue F. pratensis Huds. l. 10 Couch grass Agropyron repens (L.) P.B. l. 11 Mountain cat’s-tail Phleum montanum C. Koch. l. 12 Timothy-grass Ph. pratense L. l.s. 13 Wood club-rush Scirpus silvaticus L. l. 14 Hairy wood-rush Luzula pilosa (L.) W. l. European woodland 15 Carex silvatica Huds. l. sedge 16 Caucasian sedge C. caucasica Stev. l. 17 Red clover Trifolium pratense L. l.s. fl . 18 Fodder galega Galega orientalis Lam. l.s. 19 Spring vetchling Lathyrus vernus (L.) Bernh. l.s. 20 Meadow vetchling Lathyrus pratensis L. l.s. 21 Hogweed Heracleum sp. l.s. 22 Bloody cranesbill Geranium sanguineum L. l. 23 Wood cranesbill G. silvaticum L. l.s. 24 Caucasian cephalaria Cephalaria caucasica Litv. l. 25 Tatarian cephalaria C. gigantea (Led.) E. Bobr. l. 26 Wood avens Geum urbanum L. l.s. fl . 27 Wild angelica Angelica silvestris L. l. 28 Common oregano Origanum vulgare L. l.s. fl . 29 Hypericum Hypericum sp. l.s. fl . 30 Fireweed Chamaenerium angustifolium (L.) Scop. l.s. 114 U.A. Semenov The Wisents of Karachay-Cherkessia Table 10. (continuation) Broad-leaved 31 Epilobium montanum L. l.s. willowherb 32 Wood sorrels Oxalis acetosella L. l.s. 33 Bellfl ower Campanula sp. l.s. 34 Stinging nettle Urtica dioica L. l.s. 35 Annual nettle U. urens L. l. 36 Groundsel Senecio sp. l.s. fl . 37 Solomon’s seal Polygonatum multifl orum All. l.s. 38 Chervil Anthriscus scandicina (Web.) Manst. l.s. 39 Meadowsweet Filipendula ulmaria (L.) Max. l.s. 40 Multifl owered buttercup Ranunculus polyanthemus L. l.s. fl . 41 Lady’s mantle Alchemilla sp. l.s. fl . 42 Coltsfoot Tussilago farfara L. l. 43 Dandelion Taraxacum sp. l. fl . 44 Caucasian comfrey Symphitum caucasicum M.B. l. 45 Tall meadow cowslip Primula macrocalyx Bge. l.s. 46 White butterbur Petasites albus (L.) Gaerth. l. 47 Yellow bedstraw Galium verum L. l.s. 48 Hoary plantain Plantago media L. l. 49 Common wormwood Artemisia vulgaris L. l. 50 Fringed campion Silene multifi da (Ehrh.) Pers. l.s. 51 Ground elder Aegopodium podagraria L. l.s. American white 52 Veratrum lobelianum Bernh. l. hellebore 53 Three-lobe beggar-ticks Bidens tripartites L. l. Alpine leek (victory 54 Allium victorialis L. l.s. onion) 55 Curled thistle Cirsium crispus L. l.s. 56 Monk’s-rhubarb Rumex alpinus L. l. 57 Woodruff Galium odoratum ( L.) Scop. l. 58 Wood fern Dryopteris sp. l. 59 Male fern D. fi lix mas (L.) Schott. l. 60 Meadow horsetail Eguisetum pratense Ehrh. s. 61 Beard lichen Usnea barbata L. all 62 Bracket fungi Polyporaceae all 63 Dense-fl owered barberry Berberis densifl ora Boss et Buhse. l.br. Chapter 7. Diet 115 Table 10. (end) 64 European white birch Betula verrucosa Ehrh. br.l. 65 European spindle Euonymus europaeus L. br.l. ba. 66 Elderberry Sambucus nigra L. br.l.s. 67 Oriental beech Fagus orientalis Lipsky br.l.ba.fr. 68 Mezereon Daphne mezereum L. l. 69 Common hornbeam Carpinus betulus L. ba.l.br. 70 English oak Quercus robur sp. sh.l. 71 Blackberry Rubus hirtus Waldst. et Kit. l.s. 72 Caucasian spruce Picea orientalis L. ba. 73 Fly woodbine Lonicera xylosteum L. br.l. 74 Goat willow Salix caprea L. br.ba.l. 75 European white elm Ulmus glabra Huds. br.l.ba. 76 Guelder rose Viburnum opulus L. br.l.fr. 77 Norway maple Acer platanoides L. l.br. ba. 78 Caucasian maple A. trautvetteri Medw. l.br.ba. 79 Alder buckthorn Frangula alnus Mill. l. 80 Common hazel Corylus avellana L. br.l.ba. 81 Caucasian lime tree Tilia begoniifolia Stev. l.ba.br. 82 Raspberry Rubus idaeus L. l.s 83 Juniper Juniperus sp. ba. 84 Alder Alnus sp. br.l.ba. 85 Aspen Populus tremula L. br. ba. 86 Nordmann fi r Abies nordmanniana L. ba. 87 Mountain ash Sorbus aucuparia L. br.l.ba. 88 Caucasian currant Ribes biebersteinii Berb. s.l. 89 Scots pine Pinus hamata (St.) Sosn. ba. 90 English yew Taxus baccata L. ba. 91 Caucasian whortleberry Viburnum arctostaphylos L. l.s. 92 Bird cherry tree Padus racemosa (Lam.) Gilib. l.ba. 93 Dog rose Rosa canina L. l.sh. 94 European crab apple Malus communis L. fr.l.ba. 95 Caucasian pear Pyrus caucasica Fed. fr.br.l. 96 Caucasian rhododendron Rhododendron caucasicum Pall. l.br.

Note. Eaten parts: l — leaves; s — stems; fr — fruit; ba — bark; fl — fl owers; br — braches, sh — shoots, r — roots. 116 U.A. Semenov The Wisents of Karachay-Cherkessia Studies of the wisents’ diet in other regions of the Caucasus are of great in- terest as a potential bank of data on the possible feeding resources because the feeding composition changes not only geographically but also by season and by local dwellings (Aleksandrov, 1958; Kalugin, 1968; Kazmin et al., 1992; Litvinova, 1980, and others). The most complete data are available for the Caucasian Nature Reserve where full-scale researches of the regional feeding resources were carried out in the 60s. The resulting list contained 325 plants but, according to S.G. Kalugin (1968), that number could grow up to 400 plant species depending on extension of the animals’ dwellings and on their explora- tion of new landscapes. The scientist mentioned in his study that the wisents were very choosy about both species of the plants and their vegetative parts. For example, leaves of trees and shrubs were more nutritious and preferable for them than grass on a small pasture land, and contained up to 16% of pro- tein, 3–5% of crude fat, up to 54% of nitrogen-free extracts and 12–17% of dietary fi ber. So the reintroduced animals showed their high capability of adaptation to the local feeding conditions and high selectivity when choosing food. They eagerly eat tall-stem grasses and other plants at an early vegetation stage. Woody forage and bark of trees and shrubs play an important role in their life. Although the wisents feed on them throughout the year, they become most important during the winter period (Zablotskiy, 1957а). The experience showed a high level of similarity between the diets and energy demands of the wisent and the American bison. Although it is believed that the wisent digests the woody dietary fi ber bet- ter (Kholodova, Belousova, 1993), the share of woody forage in the diet of the bison dwelling in Alaska and grazing along river valleys reaches 94% (Camp- bell, Hinkens, 1983). During the spring-summer period, grasses, mainly gramineous and legumes, prevail in the diet of the Arkhyz population of the animals. The wisents start actively visiting salt licks at that time. Even in their early studies dedicated to the Caucasian wisent, scientists noted the high importance of the mineral nutri- tion in their life (Filatov, 1912). N.Ya. Dinnik (1897) wrote that paths from all directions led to such natural mineral sources, and that such places might be heavily hoofed as he described the salt consumption places of the wisents. Ac- cording to many scientists (Nasimovich, 1938; Bashkirov, 1939; Kalugin, 1968; Bobyr, 1992, Semenov et al., 2001), the wisents in the Caucuses visit salt licks throughout the year consuming up to 53 g of salt per day or 8 to 13 kg per year (Danilkin, 2005). As the plant associations of the North and North-West Caucuses form salt defi cient pastures, the ungulates dwelling there consume considerably more salt than the same species dwelling on the plains. For example, in the wisent nursery of the Okskiy Reserve, the average annual consumption of salt by one Chapter 7. Diet 117 wisent did not exceed 4.5 kg (Kiseleva, 1974, 1978). Chalk and salt bricks were given to the wisents as mineral feeding there. The most active salt con- sumption was registered from July till October. It is worth mentioning that the wisents ate rotten tree stumps and ant colony construction materials there in early spring. In Arhkyz the wisents satisfy their mineral feed requirements using mineral water that they drink from the orifi ces of springs and consuming the salt put into the artifi cial salt licks. During the fi rst years of reacclimatization, they were not seen at natural salt resources and consumed salt on artifi cial salt licks only in spring and summer. The wisents started actively visiting several orifi ces of mineral water springs in a number of years only when they had explored the majority of the chine and discovered those places. However, the mineral water sources are used by them unevenly. The animals preferred the water source in the Kizgych headstreams over the mineral waters in the swamp areas and on the fl oodplain probably because of the different concentration of minerals and taste. In February 1982, G.Ya. Bobyr (1992) described the case when a group

Figure 59. A Female Red Deer on the Wisents’ Artificial Salt Lick. Photo by U. Semenov with hunting camera. 118 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 60. A Herd of Male West Caucasian Turs on the Wisents’ Salt Lick. Photo by U. Semenov with hunting camera.

of 16 animals, including females and calves visited the mineral water source in the river headstreams. After the animals drank enough mineral water, they went back to the hay stacks where they had been feeding before. The next visit to the source was registered in April. Not only wisents visit the artifi cial salt licks. Deer, turs, and even bears show up there in a while. Salt is most frequently put under the trees in unexposed plac- es where the animals eat away down to 40–50 cm-deep sections between the tree roots. The wisents visit salt licks at any time of the day. The majority of peaks of the wisents’ salt consumption activity happen during the spring period when the animals start grazing on new grass and during the autumn period when changes occur in their organisms related to the biological cycles and their switching to dead grass feeding. A.A. Nasimovich (1938) noted that the spring salt consumption activity of wild ungulates coincides with the months when they more actively graze on greenery, as well as with the period of calving and milk feeding. During that period, the females feel an increased demand in mineral feeding. The salt consumption activity in autumn is associated with the rutting period of Chapter 7. Diet 119 the wisents when the feeding behavior changes both for the females in heat and for the bulls on the peak of their sexual activity with typical changes in their feeding cycles. The artifi cial mineral feeding of the wisents in Arkhyz continues. 30–40 kg of pressed salt or salt bricks are put for the animals ev- ery year. No specifi c water drinking places of the wisents were registered because the territory of the Kizgych Chine is rich in springs, creeks and small rivers fl owing down the chine bottom and forming the Kizgych River. Only great thirst can force wisents to drink from a stagnant body of water, otherwise they drink pure fl owing water. During hot summer months, the animals may drink several times a day while on cloudy days and in cold weather, they drink water once in a few days. The animals approach water as a group but keep at a distance from each other and almost never start drinking at the same moment. The animals make short pauses in their drinking process, standing in the water sometimes, and then drink again. After satisfying their thirst, some species move aside and start feeding on the branches, bark or grass available nearby. The group leaves

Figure 61. The Visited’ Salt Licks are Often Visited by Bears. Photo by U. Semenov. 120 U.A. Semenov The Wisents of Karachay-Cherkessia

Figure 62. A Male Red Deer on the Wisents’ Salt Lick. Photo by U. Semenov with hunting cam- era.

when the thirst of all of the animals is satisfi ed. The consumed water volume depends on the season, weather, age and condition of the animal. An adult wi- sent in captivity drinks up to 60 liters and sometimes up to 13 buckets of water per day (Danilkin, 2005). Chapter 8. Social Structure Dynamics and Herds Behaviour

European bison are mostly sedentary animals although some solitary species and small groups of the Arkhyz population sometimes make long-distance mi- grations. The animals live in groups of different size or solitarily, which proves the complexity of their social structure. They preferred to stay in a large herd in the very beginning of the reacclimatization period after they had been released in the Kizgych Chine. The animals got accustomed to the area by the end of the second year and started leaving the main herd in smaller groups of one to three species, or more. Thus, a group of seven species spent a year in the fl oodplain and survived the winter of 1968/1969 without any severe diffi culties. Another group of six species moved up to the Chegordala isolated terrain in early winter, came back only by late May and joined the others. A female wisent could not come through the snow drifts and broke from the group. It got so weak that the nature reserve employees had to feed it. The animals started entering the Subal- pine zone as far as the snow was melting and fi nally got familiar nearly with the entire Kizgych Valley. On August 7, the whole herd went to the neighbouring Psyzh Valley, from which it was driven back on August 20. Between January and April 1970, all the 12 European bison stayed in a single herd in the areas of the Small Chegordala and Sofi yskoe Sedlo isolated terrains. Some solitary species and small groups of two to three species started leav- ing the main herd by late April but they did not make any long-distance migra- tions. The animals moved to the Bugoy-chat Mountain Pass and lived beyond the forest border in July, and by the month end they migrated to the Cheget-chat isolated terrain, where they chose windswept areas in order to keep away from gnats and fl ies. The main herd was divided into three groups by the middle of September: the fi rst group of eight species migrated to the Sofi a isolated ter- rain, the second group of three animals stayed in the former location, while one European bison chose solitary life. The larger group came back to the former location approximately in two weeks, the second group migrated to the West of the Tomak-kaya isolated terrain, while the bull stayed separately and showed up only on October 10. All the wisents came together by the month end again and stayed in a single herd until the middle of January. Small snowfalls became more frequent by the month end, so the animals had to move toward the lower part of 122 U.A. Semenov The Wisents of Karachay-Cherkessia the chine up to the Post glade, where the herd broke in two groups again. The fi rst group consisted of several cows and a bull, while the other group included all the remaining bulls. Both groups kept close to each other between the Post glade and the Chegordala isolated terrain. Smaller groups and solitary species started breaking from the larger groups again in late April. All in all, the animals survived the winter quite well, and their condition was deemed “good”. The animals made a single herd in winter and spring of the next year and spent fi ve months together but started dividing into smaller groups and migrat- ing beyond the forest border in late June. They lived in smaller groups of two to seven species until the middle of October but their composition used to change. There were found several solitary species. The main herd was dramatically broken into several groups of two to eight spe- cies by late winter of 1971/1972, although they had kept together at the same time of the previous years. Sometimes cows with calves broke from their herds (sometimes accompanied by a bull) and lived separately. The occurrence of soli- tary bulls increased but they tended to build own groups. It is worth underlining that different groups met for a rest in the woodlands of the fl oodplain terrace and the fi r forests, built new groups of new gender and age composition and migrated to new feed areas. The animals kept breaking into smaller groups in spring, summer and au- tumn (before the rutting season). The cows also started leaving the main herd in groups of two to three species and kept a distance of several kilometres to the neighbouring groups. Like a year before, a pregnant cow left the herd and did not show up until its calf became stronger in early September. Three years later, after delivery of new animals, the herd broke into two large groups, thus, spending spring and summer separately, and came together by win- ter again. It is also worth mentioning that solitary animals and smaller groups of two to three species kept leaving the herds for a while but always came back to their large herd group. The occurrence analysis showed that the animals tended to build larger herds in winter and break into smaller groups in warm season. In the end of 1973, the herd consisted of 16 animals, including seven adult bulls, fi ve adult females, one young cow born in 1971, one young cow born in 1972, and two calves born in 1973. The observers point out that the animals seldom tended to build large herd groups in 1973. They lived in small groups in the river mouth of Kizgych in winter. Only one to two bulls permanently lived with cows. In 1974, the herd consisted of 18 animals, including seven adult bulls, six adult cows, one young cow born in 1972, two calves (a cow and a bull) born in 1973, and two calves (a cow and a calf with undefi ned gender) born in 1974. Therefore, the gender relation came close to 1:1 upon the death of two cows in 1968, which is typical gender relation of nearly all the ungulates living in optimal condi- Chapter 8. Social Structure Dynamics and Herds Behaviour 123 tions. Afterwards, despite the death of several animals the Arkhyz the herd size increased due to new born wisents and amounted to 19 species in the beginning of 1978. In spring of the same year, seven wisents (four bulls and three cows) were delivered to the Kizgych Chine from the Okskiy State Nature Reserve in order to accelerate the breeding rate and prevent inbreeding. The new animals were released to the fl oodplain terrace of the middle reaches in the area of the hay glades. Therefore, there were 29 wisents in Arkhyz (including new born and newly delivered species) in the end of 1979. The population consisted of four old bulls, 10 adult cows, eight young bulls, four young females, and three calves born in 1978 (a bull and 2 cows). The newly released wisents migrated to the upper reaches where they joint the “local” animals. The appearance of those new animals resulted in building a new herd, to which they kept for a long while until a new young bull was driven off by a “local” adult bull. The outcast bull moved to the upper reaches of the neighbour Arkhyz chine. It stayed in the Shorovo isolated terrain until October, where it sometimes grazed together with the domestic cattle. After the domestic cattle came to another place, the wisent came down to the mouth of the Psyzh River where it stayed until January 1979. It is not clear what happened to it later, perhaps it joined the herd. The animals mostly lived in the upper part of the chine in several groups of two to fi fteen species in December and the fi rst half of January. The bulls lived separately from the cows and young species. Later on, the animals migrated to the lower part of the chine, and between May and the mid-autumn lived in the Subalpine and Alpine meadows coming up to the altitude of 2,600 m above sea level in the area of the Sofi yskoe Sedlo Mountain Pass. Moreover, they never stayed in the Alpine zone overnight and always went to the forest. In the morn- ing the wisents came back to the upper border of the Alpine meadows for feed- ing, stayed there until the evening choosing windswept areas in order to keep away from gnats and fl ies, and then came back down. In the warm season, the altitude range of their migrations and the duration of their stay in the open areas mostly depended on the weather conditions. On sunny days, they came higher and stayed there longer than on dull days. They browsed and rested next to the upper border of the forest and Subalpine zone in case of nasty weather. The herd structure and group size also depended on the season. The wisents lived in three groups consisting of 23, 4, and 2 species, respectively, in the winter of 1979/1980. The fi rst group was built by the wisents of different gender and age, including the calves born in 1979. Two other groups consisted of four adult bulls and two (a young and an adult) bulls that lived outside the nature reserve in the Psyzh isolated terrain. The population started breaking into smaller groups 124 U.A. Semenov The Wisents of Karachay-Cherkessia and solitary species in spring. Such social structure remained until late August. The herd got larger in the beginning of the rutting season. The yearly average gregarious ratio amounted to 8.0 in 1980. Between February and March 1982, the herd was divided into three more or less permanent groups: a group of 19–23 male and female animals of different age (sometimes, one to four species left the group but came back in one or two days), a group of four adult bulls and a group of an adult and a young bulls. The animals built four groups in late May. A young bull joined the third group, the group of four bulls remained unchanged, seven cows of different age, two calves born a year before and an adult bull kept together, while the fourth group consisted of 12 male and female animals of different age. The next year the recorded gender and age structure of the population looked as follows: 11 bulls and 14 cows aged over one year, four adult animals (gender not defi ned), and four calves born in 1983 (gender not defi ned). Calves were brought by the females aged six to seven years that year. The wisents showed up in groups of 2 to 18 male and female animals of different age in winter and spring. The young cows that gave birth to calves that year, as well as the youth aged over one year made up a group of 12 species and lived together for about 10 months. The size of the other groups changed. The bulls tended to form small and medium groups of two to six species. Sometimes there were found solitary animals that kept away from the herds. The group building criteria (gender and age) depended on the periods in the life of the animals (rutting season, snow season, birth of calves, etc.). As a rule, large herds are built during the rutting season or in case of thick snow cover, but sometimes it happens at random due to association of several groups, which is related to the search for the most favourable feed and rest locations. In such a case, the associated herd is very instable and is used to break after a while, thus, building new groups, sometimes with a different composition. For example, a group of eight adult cows and an adult bull joined a group of seven adult bulls in the Chegordala isolated terrain on March 12, 1983. Together they migrated to the area of the Grushevaya glade, stayed there until March 18, then broke into sev- eral groups and left the area. In autumn the main population consisted of three groups of 15, 12 and 3 species, respectively. The fi rst two groups consisted of cows, calves and bulls, while the third group consisted of adult bulls. Some bulls lived away from the herds but sometimes joined one group or the other. By the middle of the 1980-s when the wisent population increased by more than two times and amounted to about 35–37 animals, their social structure changed as follows: as the winter set in, and the snow cover reached more than 40-50 cm the wisents gradually made one herd and kept to it until late February or even late March. Afterwards, it was divided into several smaller herds belonging to a certain family, gender or age groups. The strongest associations were built by cows with calves of the current year and amounted to 10–20 animals. Their herds Chapter 8. Social Structure Dynamics and Herds Behaviour 125 grew up to 32–35 animals if joined by young bulls. It is worth mentioning that the social structure of the Arkhyz wisent population is variable and depends on the season and the total amount of species. The population growth by means of the natural increase resulted in the changes in the herd groups. While the animals seldom left the main herd in smaller groups, and the group size mostly amounted to one to three species in the fi rst years of reacclimatization, in 15 years it was common to build groups of 4, 5, 7, 10, 15, and over 20 species in spring and summer. In the beginning the groups were built at random, however, afterwards, they were more gender or age oriented and sometimes based on family relations. For example, in the winter of 1981, the observers found several gender and size oriented groups that survived until the early spring of the next year. The largest herd had 19 animals and from time to time was joined by some solitary bulls or groups of two to four bulls. The social structure changed next spring. The main herd was broken into several groups of 3, 4, 10, and 12 species. The fi rst two groups consisted of adult bulls. The third group consisted of seven cows, two calves born a year before, and an adult bull. The fourth group was the largest and consisted of male and female animals of different age (Bobyr, 1992). The observers found relatively large herds of up to 32 animals in the snowy winter of 1985/1986. Such herds mostly consisted of cows, calves aged below one year, and young wisents of up to three years. From time to time, those groups were joined by adult bulls, which broke from the herd in a while, lived soli- tarily and sometimes joined other groups. They also found some solitary bulls, small groups of two to three bulls, or cows with young wisents. The gender and age structure of the recorded wisents looked as follows: 18 adult bulls, 22 adult cows, 8 young wisents aged up to three years (gender not defi ned), and 6 calves of the current year. The wisents again built a large mixed herd of 37 species and several smaller groups in March 1988. However, in a while the main herd was divided into three herds of 6, 15, and 16 species. Two bulls kept living separately outside the nature reserve. The wisents as usual built a large herd of 39 species and some smaller groups of three to fi ve species in the late autumn and early winter of 1988/1989. The herd fell apart into several groups based on the family, and gender and age crite- ria in spring. In late March, the observers found a group of 16 species (mostly fe- males) in the lower reaches not far from the avalanching. The group consisted of three young wisents aged two or three years and a calf born a year before. They also found two more groups of eight species (one group also had two calves born a year before) upstream in the avalanching area. Adult and young bulls built smaller groups of three to fi ve species, sometimes kept separately from the other herds, then joined some other group and left in a while. Next autumn and winter, a group of 12 wisents (mostly females and three calves of the current year) were found in the Byky isolated terrain. The animals spent 126 U.A. Semenov The Wisents of Karachay-Cherkessia the entire year in that area making small migrations of 1–3 km and coming back to the former location. A group living in the area of the Krest glade consisted of three adult bulls, nine adult cows, three calves born a year before, and a young wisent aged 2–3 years. The third group inhabiting the Chegordala isolated ter- rain consisted of 19 species, including four bulls, eight cows, six young wisents, and a calf born that year. That group had the smallest amount of the calves of the current year. Probably, there were too many old females with the interbirth interval of over 3 years. The Arkhyz wisents used to build medium-size groups of 10–15 species in the beginning of 1990s. The observers found such groups most often, while the frequency of large herds of 25–30 species grew shorter. Solitary species and groups below eight to nine species appeared more seldom. Due to the deep snow cover, the animals moved from the open areas to the forests where they built larger herds. The observers found a herd of 29 species (mostly females) in the shade-tol- erant temperate coniferous forest on January 17, 1992. The herd consisted of 17 cows, fi ve bulls, four calves of the current year and young species aged 2–3 years (gender not defi ned). The calf gender relation could not be defi ned. How- ever, their total number registered within the previous years proved low birth and population growth rates. The gender structure analysis showed that the adult spe- cies prevailed in different herds. Low specifi c weight of young species might be caused by signifi cant amount of old infertile cows, or depression of the animals due to the increasing affi nity breeding. The next year the observers found some changes in the gregarious behaviour. The wisents mostly lived in groups of 2–18 male and female species of different age. As a rule, they consisted of cows, young wisents aged three of four years, calves of the current year, and one or several young bulls. Several groups came together and then left again in the early winter and autumn. The analysis of the gender and age relation of the recorded groups of all in all 47 species showed that the groups mostly consisted of 21 cows, 17 bulls, seven young wisents aged two or three (gender not defi ned), and two calves of the current year. The similar social structure trend remained in 1994, when the Arkhyz population consisted of three more or less permanent multi-gender and multi-age associations. The herd inhabiting the Baga-tala isolated terrain consisted of six adult bulls, two females with calves of the current year, fi ve young wisents aged over two years, and 11 multi-age species (gender not defi ned). The Byky isolated terrain was inhabited by three adult bulls, three young species aged two or three years, and six adult cows without calves. The observers failed to defi ne the social structure of the group inhabiting the Tokmak-kaya isolated terrain. According to the Wisent Register of the Teberda State Nature Reserve as of February 1, 1997, the Kirgych Mountain Valley was inhabited by fi ve wisent Chapter 8. Social Structure Dynamics and Herds Behaviour 127 groups. One of them was found in the Eki-Su-Arasy isolated terrain. It con- sisted of three adult cows and three calves born a year before. The second group dwelled in the Tokmak-kaya isolated terrain and consisted of 16 males and fe- males of different age. The group found in the Geleu-tala isolated terrain con- sisted of two females and three bulls, while the group dwelling in the Chegordala isolated terrain consisted of four bulls. The last group inhabited the Byky-ayagy isolated terrain and consisted of seven adult cows and fi ve bulls. The calves of the current year amounted to 6.9% in the total amount of the recorded animals. There were four separate groups recorded in the area of Kizgych in 1998. The fi rst group of 14 species sometimes was found not far from Tokmak-kaya and consisted of six adult cows, four adult bulls, and four young wisents (gender not defi ned). The neighbouring herd of nine species consisted of fi ve adults and four youths aged two to three years. In is worth mentioning that other groups did not have any calves of the current year. The third group consisted of four adult females and three young species aged two years. The fourth group of fi ve females and four bulls sometimes broke into two separate groups of bulls and cows that lived separately for a while and then came together again. In 1999, the four herds remained but their structure changed. The popula- tion of the herd inhabiting the area of Tokmak-kaya decreased and amounted to 10 species (fi ve adult cows, two young cows, and three adult bulls). Perhaps, the gender structure changed due to more precise gender defi nition as previously the observers failed to defi ne the gender of some species. Still, it did not signifi - cantly infl uence the pattern of the herd structure. In the late 1990s, the number of the Arkhyz wisent population fl uctuated be- tween 38 species in 1996 and 46 species in 1999. The population recording car- ried out in spring 2000 showed that the total amount of the animals was 20– 21 species. In the course of the Kizgych Chine examination the observers found four groups: fi ve species in the area of the water fall not far from the Volchye Mesto draw, eight to nine species in the Byky isolated terrain, four adults in the upper reaches of the Kizgych River, as well as three bulls (two adult ones and a young one) at the avalanche cone. Later they found ichnofossils (litter of young animals and old traces) of another group with the estimated population of up to four species that was not recorded1. The population decrease changed the social structure. The animals started building smaller groups and herds. The animals’ and ichnofossils’ occurrence analysis in 2000–2003 showed general reduction of the species in the groups. The diagram (Fig. 60) shows that the solitary species amounted to 12% of the population. Most of the solitary species (70–80%) were adult bulls. The occur- 1 Such data needed additional investigation and were not included into the consolidated report — author’s note. 128 U.A. Semenov The Wisents of Karachay-Cherkessia rence of solitary bulls increased in August–September due to the rutting season and replacement of old and young bulls by stronger species. As a rule, upon the rutting season and activity reduction, the outcast species joined some groups when the animal antagonism was not so strong. Having spent a while with other animals, old bulls started leaving the groups and fi nally got entirely isolated from the main herds. After a while they were found dead. Old infertile cows sometimes also choose solitary life. They live alone for a while, and then get weak and vulnerable to the predators or diseases. However, not only old animals live solitarily, sometimes, even young wisents do not join the main groups. Closer examination showed that they were hidebound and nearly did not make any long-distance migrations. Their migration radius did not exceed 1 km. All of them eventually died within three to nine months. Thus, in the beginning of the 1980s, the observers found a dead cow that had previously lived separately from the other groups. It was very hidebound. The thanatopsy showed a brain tumour, and stomach and intestine infl ammation (Bobyr, 1992). Between 1977 and 1985, there were found some solitary animals that were ill and hidebound, they limped and coughed. All such animals died within several months. The analyses of solitary wisent occurrence shows that the Kizgych wisents are not used to solitary life, only mature and healthy bulls can live alone for a short while and then join the groups. The healthy females break from the herd only during the calving season, though old and ill cows (aged aver 16 years) also leave the herds (Nemtsev and others, 2003), though some authors have evidences that cows may remain fertile until the age of 25 years (Sokolov, Tembotov, 1993). The next social structure type includes groups of two to three species (occur- rence amounts to approximately 23%). Such groups occurred more seldom dur- ing the fi rst years of reacclimatization and did not occur at all in snowy winters. In such periods, they built larger groups and relatively large herds of 15–20 spe- cies. Small groups can be built by male and female animals of different age. They may consist of a female with a calf, a barren female (both old and young), several females and bulls of different age (not only during the rutting season). Such groups as temporary associations are very instable. It is worth mentioning that the occurrence of small groups was increasing since 2000, probably, due to the general wisent population reduction in the Teberda State Nature Reserve. Groups of four to fi ve species appeared to be most common (34%). Their occurrence increased in the last years. As a rule, they consist of a cow with a calf born a year before or that year, a barren female and a bull. Sometimes they were joined by old females or young bulls, but the presence of the latter changed within the year. Bulls used to stay with their females during the rutting season but afterwards could easily leave the group and join another one. The gender behaviour of the bulls changes during the rutting season when they are forced to Chapter 8. Social Structure Dynamics and Herds Behaviour 129

Figure 63. Gregarious Behaviour of the Wisents in the Teberda State Nature Reserve (be- tween 2000 and 2003). join the groups but by the end of that season the motivation of loyalty expires, so the bulls get more tolerant to their opponents and are able to build larger groups. The size of the groups of bulls of different age is very instable. Solitary species or groups of two species leave the main group, live separately for a while and then join a mixed group. Male groups lack loyalty to a certain group; bulls keep together as long as there are no reasons for pronounced competition during the rutting season and as in a group it is easier to keep away the predators. Having joined old bulls, young bulls learn how to survive outside their mothers’ herds. A most signifi cant component of communicative behaviour is the ability of a young bull to join an adult bull or a male group. The stronger and more expe- rienced the “tutor” is, the more effi cient the adaptation of a young bull to the independent future life is. Groups of cows and young species are more stable. Moreover, females with calves of the current year left very seldom and joined other groups. In comparison to bulls, the female associations were often caused by “family” relations. Those animals built the core of a group and were joined by other animals. Increased group size and herd building (up to 9–10 species with the occur- rence of 28%) happened by late summer as a cause of the beginning rutting season. This is the time when the animal behaviour changes, and bulls get more irritable and aggressive. Fighting for the cows, the stronger bulls oust younger and weaker species and can even break the main group into several smaller ones. Therefore, such associations are very instable and sometimes fall apart into sev- eral groups that come together very often. As the rutting season comes to an end, the animals become calmer, migrate more consciously, and build more dynamic groups. Such mating groups can be signifi cant. For example, such groups have a 130 U.A. Semenov The Wisents of Karachay-Cherkessia size of up to 97 species in the Caucasian Nature Reserve (Nemtsev et al., 2003). The Arkhyz wisents built the largest groups of up to 19–20 species in August- September, however, recently, such associations were seldom, and their occur- rence amounted to 3%. The wisents built three separate groups in spring and summer 2001. The ob- servers found two groups of four and eight species in the upper part of the val- ley, i.e. in the Baga-tala and Bugoy-chat isolated terrains, respectively. The fi rst group consisted of adult bulls, and the second one — of males and females of different age, including young species. The animals did not come together de- spite their proximity and migration between the same stations. The third group of six species, including the youths inhabited the Byky isolated terrain. In early autumn, the animals came together for a while, and then the herd fell apart into smaller groups that fed and rested separately. Four adult bulls built a group and lived in the Baga-tala isolated terrain. Seven males and females of dif- ferent age inhabited the same area, the fl oodplain terrace of the right bank. Two groups of two species each lived downstream in the Byky and Tokmak-kaya isolated terrains. The fi rst group consisted of two bulls, while the gender and age structure of the second group was not defi ned. The calves of the current year were found in neither of those groups. The herd structure and group size remained nearly the same the next winter. Within a year, the animals migrated in smaller groups between the main stations. Three adult bulls stayed in the Tokmak-kaya isolated terrain, and four adults (gender not defi ned) inhabited the mountain pass in the Byky-ayagy area. Later on, the observers found an adult cow with a young species next to the fi rst group. All in all, there were found 11 animals, including seven adult bulls, an adult cow, a young cow, and two more species (gender not defi ned). There were also found traces of one more wisent group of four to fi ve species downstream. The calves of the current year were not found. The same structure remained in 2005. Adult bulls built a group of four species and stayed in the Tokmak-kaya isolated terrain without making any long-dis- tance migrations or breaking into smaller groups. Two other groups of two spe- cies each lived midstream, sometimes came together for a few days, then they came apart for a while. There were also found some solitary species in that area, probably they broke from the above groups. The previously found cow and young bull stayed in the Byky-ayagy isolated terrain, while the mixed group of fi ve species migrated closer to the Baga-tala isolated terrain and during all the seasons stayed there making only short-distance migrations to the Subalpine and Alpine zones. No cows gave birth in 2005/2006, so there were no calves of those years in the groups. Within the next few years, the wisent spread in the Kizgych Chine remained more or less the same. The animals moved in small groups without building Chapter 8. Social Structure Dynamics and Herds Behaviour 131 herds even in autumn. Signifi cant population degeneracy and decrease signifi - cantly changed the gender behaviour of the animals. Thus, the wisents did not build any mating associations. New-born calves were not found within several years. Only in November 2007, there was found a group of fi ve species, includ- ing a wisent aged one-and-a-half years in the Post isolated terrain. Moreover, that year there were found traces of some adult species, as well as a young spe- cies next to the salt spring of Baga-tala. Probably, those were the traces of the previously found group as afterwards no young species were found. The population structure did not change in 2008 except for the group of four bulls that broke into two smaller groups of two species each and inhabited the Tokmak-kaya isolated terrain. They lived in the same area, but fed and rested separately. Other animals continued living in small groups all over the Kizgych Valley. Two groups of two species each were found midstream, and a mixed group of four to fi ve species was found not far from the Baga-tala isolated ter- rain. There were found no new-born calves that year, as well. In 2009, the social structure changed a bit: one of the male groups of the Tokmak-kaya isolated terrain fell apart, and a bull joined another group of two adult bulls. Nothing is known about the further fate of the other bull. Two groups of two species each stayed midstream. The observers failed to defi ne the gender of those species but their behaviour over the year and traces let them suppose that those were bulls. A group of fi ve to six species was found midstream in late April. The group stayed there until October. The traces of a new-born calf of that group were found in autumn. A young bull left a small group of three species in April 2010. The two re- maining adult bulls stayed in the area of Tokmak-kaya. Two other groups of two species each kept apart within the fl oodplain midstream area of the Kizgych River. The observers found traces of two adults in the neighbourhood of Baga- tala. A large group of three to four wisents, including the previously found calf inhabited the upper reaches. Only fi ve wisents inhabited the Arkhyz area of the Teberda State Nature Re- serve in 2012. Two adult bulls, an adult cow and a young cow aged 2 years built a group and stayed for a while in the area of the spurs of Sofi yskoe Sedlo Mountain. An old solitary bull also lived in that area and sometimes joined the group. The decreased population resulted in the animal concentration in the area; the animals nearly did not leave the nature reserve. Moreover, they inhabited the upper reaches of the Kizgych River where they stayed during all the seasons. Chapter 9. The Wisent Reproduction and Calving Seasons

According to the earlier studies of the native Caucasian wisent (Vinogradov, 1870, 1871; Dinnik, 1910; Filatov, 1910, 1912), the rutting season started in August and ended in middle of September. N.Ya. Dinnik (1910) underlined that the bulls hunted at that time were hidebound and had empty stomachs. Agitated bulls could not keep quiet, hoofed the earth, rubbed with their neck, muzzles and horns against young trees in the stations, etc. According to A.F. Vinogradov (1870), the rutting season of the Caucasian wisents lasted only for about three weeks since late August and until the second decade of September. However, in his study of calf capture within the Kuban Hunt of the Grand Duke, Dinnik (1910) insisted that on April 10 there were found calves born a few days before. The capture lasted until May 21, and the calves found last had also been born a few days before. As the cow pregnancy lasted for about nine months, and new- born calves were found both in the fi rst decade of April and the third decade of May, the rutting season of the native Caucasus wisents lasted for approximately two months since August and until September. The wisents of the Bialowieza Forest had the rutting season at the same time as the Caucasian wisents – in Au- gust and September (Dalmatov, 1849; Kartsov, 1903). A.A. Danilkin (2005) believes that the females can have several estrous cycles (up to fi ve) within a year, each lasts from one or two up to three or eight days. If within such period the female does not mate with a bull, it will have another estrous cycle in 9–10 days, more often in 20–30 days. Therefore, such a feature might result in a shift of the rutting season of some wisent populations. Thus, the rutting season of the wisents in the Bialowieza Forest lasts from June until August–October, sometimes even November (Kulagin, 1919, 1940; Serzhanin, 1955), in Poland — from August until September (Jaczewski, 1958), in Ukraine (Volyn) — from October until March (Tatarinov, Dyakun, 1969), in the Okskiy Wisent Nursery — since the second half of July until March–April (Kiseleva, 1973, 1978), in the Priokskiy Terrace Wisent Nursery — since July until January (Pererva, 1981), in the Caucasian Nursery — since July until January (data by Nemtsev, 1985) or since the end of July and until October, sometimes in February (data by Kalugin, 1958, 1968). As the wisents brought to Arkhyz in 1968 were too young for the rutting, the fi rst calf was born only in July 1971, though the fi rst estrous cycle was found Chapter 9. The Wisent Reproduction and Calving Seasons 133 in the last decade of October of the next year. That year the rutting features occurred until the middle of December. The table shows that the main rut- ting season lasts from September until October despite of the variable rutting terms. In the beginning of the acclimatization, such a shift might be related to the migration from different climate zones and adaptation to the new mountain conditions. The rutting season of the Arkhyz wisent population signifi cantly decreased in 10 years (in 1978): it started in late August or early September end nearly completely ended by the middle of October. The sexual activity decreased by the beginning of cold season. In 1979, the rutting season fea- tures occurred signifi cantly later (in the end of September) and lasted until late November, which, perhaps, was related to the release of the new animals in the Kizgych Valley in order to accelerate the reproduction rate, prevent in- breeding and increase the population. The rutting season shifted even in the Central Wisent Nursery upon release of the new animals. That year there were two stages of the rutting season: stage one lasted from July until September, and stage two — from November until January. Similar processes were found among the animals of the Bialowieza Forest (Kulagin, 1919), Askania-Nova

Figure 64. Lowland European Wisent Cows and Calves Aged Two to Three Weeks in the Prio- ksko-Terrasniy Nature Reserve. Photo by N. Treboganova. 134 U.A. Semenov The Wisents of Karachay-Cherkessia (Zabolotskiy, 1957) and the Caucasian Nursery in the course of reintroduction (Kalugin, 1968; Nemtsev, 1988). Table 11 Rutting Seasons of the Arkhyz Wisents (According to the Teberda Reserve Nature Records)

Years Beginning of the Rutting Season End of the Rutting Season 1972 Last Decade of October Middle of December 1973 September October 1974 Middle of August End of December 1977 End of August September 1979 End of September November 1980 September September 1985 Third Decade of September October 1986 First Decade of September Not defi ned 1987 End of August Not defi ned 1988 Middle of August Not defi ned 1989 Middle of August Not defi ned 1992 Third Decade of August By the Beginning of October 1993 September October 1994 Not Available Not Available 1995 Not Available Not Available 1996 End of August Beginning of October 1997 Not Available Not Available 1998 September October 1999 Not Available Not Available

Thus, the initially shifted dates of the beginning and end of the rutting season of the Arkhyz wisents slowly decreased in the course of the adaptation. The rut- ting season got more or less distinct frames in four to fi ve years upon the release (Table 11). Such season lasted for two months at the fi rst stage (since the middle of August until late October), while it started in late August – early September and ended in late September – early October in the late 1980s. A.S. Nemtsev (1988) believes that the rutting season and pregnancy period of the restored mountain wisents of the Caucasian Nature Reserve decreased upon the enforced naturalization. The calves were born in the beginning of the second decade of June, and standard deviation from the average birth date amounted about 30 days between 1956 and 1959. That value had signifi cantly decreased by 1985. Most of the calves were born in May (on May 21 at the average), while the deviation from the average birth date dropped from 30 down to 23 days. The native Cau- casian wisents of the 19th and 20th centuries had the rutting season approximately at the same period of August–September (Dinnik, 1910). Chapter 9. The Wisent Reproduction and Calving Seasons 135 The rutting season of the captive animals varies and might occur nearly in any time of the year. The main estrous cycle of the American bison (Bison bison) lasts from July until October; however, some mating cases were registered even in April. Some researchers believe that prolonged rutting period of the wisents is caused by the long-term captive breeding in the past when males and females were accustomed to living together all the time. They often concentrate on some areas in the nurseries (like creep feeding areas, artifi cial salt springs, watering holes, etc.), which results in estrous cycles destabilization and might provoke the beginning of the rutting season even in winter. Permanent neighbourhood of fe- male and male species reduces the competition among the animals: even young captive bulls participate in the rutting, while adult bulls would not allow it in the wild (Danilkin, 2005). According to different data sources, the pregnancy period of the wisents lasts from 265 up to 297 days. This period of the wisents in the Bialowieza Forest amounts to 257–272 days, 265 days on the average (Korochkina, 1971), in the Caucasian Nursery 261–283 days (Kalugin, 1968), the Okskiy Wisent Nursery 217–297 days (Kiseleva, 1974, 1978). Some wisent populations sometimes show signifi cant deviations from the average calf birth date, which may exceed 50–75 days. No special researches have been carried out in the Teberda Natu- ral Reserve but the occurrence analysis of the wisents with calves in May-June shows that they are quite close to the ones of the Caucasian Nature Reserve (as species living in similar environmental conditions) and amount to about 270 days. It is also proved by the shifted calf birth periods in 1980 caused by the shifted rutting season due to the release of new animals. That year some cows mated with bulls only in November, while a new-born calf was found in the beginning of August. A calf aged a few days was found in the course of the fi r forest examination in early May of 1973, repeated examination allowed to fi nd another calf born in early June. Two calves aged a few days were found in the fi r forests at the altitude of 1,350–1,500 m above sea level in May 1974. Three calves were born next year; in May, a calf aged a few days was found in the mixed coniferous forest with the Nordmann fi r prevailing; another one was found in June, and the third one — only in August within a group resting at a forest glade. The next occurrence of the calves of the current year was regis- tered in 1979: two calves (age not defi ned) were found not far from the conifer- ous forest range in June and July. The latest birth date of the calves was August 1980, when the observers found a calf aged a few days in the fi r forest at the Southern slope exposure. However, it was the only signifi cant deviation from the standard birth date of the Arkhyz calves due to the new animals’ release in the Kizgych Chine. Other calves were born between May and late June of the 1970s (Table 12). Therefore, by the middle of 1980s most of the calves were born in May, sometimes in early June. 136 U.A. Semenov The Wisents of Karachay-Cherkessia According to the earlier data (Dinnik, 1910; Filatov, 1910), none has seen any Caucasian wisent tweens. Moreover, new calves were born not every year. It was believed that cows got pregnant once in two or three years. According to the examinations in the Caucasian Nature Reserve, the female fertility amounted to 50% and in some years even to 100% upon the new animals’ release. The in- fertile cows amounted to 25–30% but that indicator decreased to 12–18% after cold winters (Nemtsev et al., 2003). There was found an interesting pattern as follows: the cows got pregnant every year in case of low density and once in two or three years in case of high density (Kalugin, 1968). No twins have been found within the entire period of observation of the Arkhyz wisent population.

Table 12 New-Born Calves Found in the Kizgych Valley of Arkhyz (According to the Teberda Reserve Nature Records)

Years Amount Date of Identifi cation Place of Birth 1- Early May 1973 2 Fir forests, at the age of a few days 1- June 1974 2 May Fir forests 1975 2 May-June Mixed coniferous forest with the fi r prevailing 1 May 27 Born in the 72nd district of the Kizgych Chine 1978 (found in the glade of Baga-tala) 2 June Not defi ned 1- June 1979 2 Shade-tolerant temperate coniferous forest 1 - July Born in blocks 73-83 of the shade-tolerant 2- July 1980 3 temperate coniferous forest and glades 1- Late August Shade-tolerant temperate coniferous forest Found in a group and migrated together with 1986 3 June adults Found in a group and migrated together with 1987 3 Third Decade of June adults Found in a group and migrated together with 1988 4 Second Decade of June adults Found in a group and migrated together with 1989 3 Second Decade of June adults Found in a group and migrated together with 1991 4 June adults The observers found two calves with females 1992 4 June keeping separately from the herds Two more calves found in autumn Found in the thick bird cherry tree woods. Two calves aged one to fi ve days were found 1993 2 May 29 with their mothers in the shade-tolerant temperate coniferous forest. Chapter 9. The Wisent Reproduction and Calving Seasons 137 The increased or decreased pregnancy periods might be caused by individual features of females (age, state, experience, etc.). The pregnancy periods of the Caucasian wisents and the Lowland European bison are nearly the same, it is believed that it lasts for about nine months (Heptner et al., 1961). As a rule, the calf lactation lasts for up to 8–10 months and depends on in- dividual features of the animals. Some cows nurse their calves even in excess of that period. According to the data of the Arkhyz observers as of 1980, the females nursed the calves born a year before until late June. It was found out that the calves were nursing for about 10–12 minutes in the beginning of the lactation period of the previous year, while at the age of 11–12 months they were nursing for 3–4 minutes in the end of such period. Probably, the new environmental conditions (mountain areas) and young age of the animals resulted in restrained fertility rate among the released cows (the fer- tility rate of the fi rst fi ve years amounted to 6%), while the parturition frequency depended on the individual features of each female. According to the researches, only 78% of the fertile cows in the Bialowieza Forest gave birth to calves, while the average parturition frequency amounted to one year and three months before the beginning of the 1980s (Korochkina, 1971; Bunevich, Kochko, 1988). In Po- land that indicator varied from 16% up to 86% and depended on the acorn crop during the year before the parturition (Bunevich, 1999; Kozlo, 2001). Within the fi rst years upon the release, the fertility rate of the Lowland Eu- ropean wisent females reached 100% in herds, though it amounted to 25–30%

Figure 65. Barren Female Arkhyz Wisents within Different Observation Periods. 138 U.A. Semenov The Wisents of Karachay-Cherkessia on the average and decreased to 12–18% after long and cold winters (Kalugin, 1968; Nemtsev et al., 2003). The fertility rate of the captive sexually mature cows of the Okskiy Nature Reserve amounted to 75–100%, while most of the cows got pregnant every year (Kiseleva, 197 3). In the course of the observations carried out in the Prioksko-Terrasniy Nature Reserve, it was found out that the fertility rate of a herd depends on the bull change. When bulls and cows were kept together for a long while, the fertility potential signifi cantly decreased and in the fi rst years amounted to 67–62%, afterward reaching 38%. The observers found out that it was required to change the bulls in order to increase captive female fertility like it happened in the wild (Pererva, 1981; Petrov, 2003). The fertility of the Arkhyz cows changed depending on the period. It used to be very low in the beginning, afterwards that indicator increased by 43% and amounted to 74 % on the average between 1973 and 1997. However, the fertil- ity rate dropped between 1998 and 2012, and the rate of barren cows in herds reached 95–98% and amounted to 97% on the average, while there were no calves born at all in some years (Fig. 65). Such conclusions shall be deemed rela- tive as some calves might have died without being found. E. Tsibizova (Tsibi- zova, 2009) underlined that the loss of calves within the fi rst two days upon their birth in the Okskiy Nature Reserve reached 52%. It is harder to fi nd dead calves in the wild as the predators fi nd them more quickly. No dead calves have been found within the entire observation period, except for the one found in the womb of its mother. Chapter 10. Moulting

The winter coat of the wisents is very solid. Therefore, the animals often lie on the snow without digging it. Their summer coat is signifi cantly shorter and not so thick. It is believed that the wisents moult once a year. It is said in the earlier studies that the Caucasian wisents started moulting in late spring and ended in late summer (Satunin, 1898, 1914; Filatov, 1912). Having observed summer fur moulting, some researchers believe that the animals moult twice a year — in spring and autumn (N. Treboganova, personal communication). De- scribing the appearance and seasonal moulting of the wisents (B. b. caucasicus), N.Ya. Dinnik (1910) pointed out that the main coat colour becomes more or less uniform in autumn, chestnut brown all over the body. Moreover, the head gets slightly darker than the rest of the body, while the fur of the beard and foreleg knees almost becomes black. The coat along the spine and withers is light chest- nut, while the long and harsh tail tip fur is black. The hair gets longer, and the undercoat gets shaggier in late August. The coat on the head gets signifi cantly longer than on the rest of the body in September. “Although the wisents’ coat is still short in September, they already have a tender undercoat that is so shaggy that it is impossible to see the skin”, the researcher underlined (Dinnik, 1910, p. 139). In winter the coat of the native wisents got dark brown and very large, especially on the bulls’ heads, where it got “curly” or “waived”. At that time, the adult wisents had a shaggy long beard on their chin and under the lower jaw near the neck, and a coarse mane from nape to withers that stuck up and made the animals look even more powerful and threatening. Long hair also covered their lower neck and breast. The female coat had the same pattern as that of the bulls, though it was shorter. “All the wisent bulls have larger mane and longer hair in the fi rst half of May. At that time, the cows also have similar winter coat, which starts moulting rapidly in May. The wisents shed so much fur in June that they seem to be nearly naked from a distance. The bulls do not have any mane at that time, though they still have long hair on the lower neck, chin and partly on the cheeks”, Dinnik pointed out describing the spring moulting (1910, p. 140). Kalugin (1958) believed that the wisents of the Caucasian Nature Reserve had moulting in the second half of February. Some species, mostly bulls, started moulting already in the second decade of February and had their fur coats com- pletely changed by the end of May. Adult cows, especially the ones with calves, 140 U.A. Semenov The Wisents of Karachay-Cherkessia moult later; moreover, some of them keep moulting until late July. The begin- ning and duration of the moulting period mostly depend on individual features of an animal: age, gender, and state. Healthy adult animals start moulting earlier and get new fur coat within a shorter period of time. Every adult wisent loses about 4–5 kg of hairs (Nemtsev et al., 2003). Small winter fur fl ocks can be found at tree and bush branches on the wisent migration routes. The moulting period of the Arkhyz wisents is long and depends both on indi- vidual features of the animals and on the weather conditions throughout the year. If spring is late and cold the moulting period starts in May, though ordinarily it begins much earlier. Within the fi rst years of reacclimatization, the wisents moulted when it was warmer and the risk of temperature drop and colds was insignifi cant; it also used to be more intense. At that time, the moulting period started in late April and lasted until early July (Table 13).

Table 13 Moulting of the Arkhyz Wisents (According to the Teberda Reserve Nature Records)

Moulting Duration, Years Beginning of the Moulting End of the Moulting Months Middle of the third decade of End of the fi rst decade of 1972 1.5 April June 1973 Early April Late May 2 1974 May Middle of June 1.5 1975 Middle of May Last decade of June 1.5 End of the second decade of 1977 Late June 2.5 April 1978 Third decade of April Middle of June 2 1979 Late April Middle of June 2 1980 Early May Late June 2 1983 Third decade of March Late June 3 1985 March Late June 4 1986 April Early July 3.5 1987 April Middle of July 4 1988 April Middle of July 4 1989 April Late June 3 1992 Third decade of February Early May 3.5 1993 April Middle of July 3.5 1994 Third decade of March June 3.5 Chapter 10. Moulting 141 The observations of 2000/2007 showed that in case of warm spring adult bulls started moulting already in late February (more often in March), and had nearly all of their winter fur shed by the middle of May. The cows with calves of the current year moulted until the middle of July. For example, in 2001, the moulting started in the third decade of March, and there was found a group of four adult bulls with summer coat on May 18. The cows moulted for a longer while that year. Several species that only started moulting were found in the spurs of the Sofi yskoe Sedlo Mountain Pass in late April. They had moulted fur fl ocks of a lighter colour on their necks. Still, nearly all the wisents inhabiting the upper part of the Kizgych Valley had winter coat already by the end of September. There was found a young bull with uncompleted moulting in the fl oodplain area in June 2003. It had old fur on the back and sacrum. The animal was very hidebound and looked ill, nothing else is known of the further fate of that animal. Upon comparison of our observation results with the Teberda Reserve Nature Records and the data of other researchers, it is worth underlining that the moult- ing periods and sequence of the Arkhyz wisents became similar to those of the native Caucasian wisents at the end of 1990s. The winter fur coat sheds fi rst from the neck, then from the forepart of the body and sides. Afterwards, the fur falls from the back, hindquarters, belly and only then — from the tail. It is noteworthy that in the beginning of the reacclimatization the animals moulted nearly until mid-May, that process got longer and more even though in a while. In the course of the generation change, the animals got accustomed to the mountain environment of the Caucasus. The moulting period of the Arkhyz wisents is a bit shifted despite the similarities of the environmental conditions of the Caucasian and Teberda Nature Reserves. That trend is most distinctive among the adult bulls. In the Caucasian Nature Reserve, they start moulting in the second decade of February (Kalugin, 1968), while in the Kizgych Valley the fi rst signs of moulting appear only in late February or early March even in most favourable weather conditions. That trend occurs also among other age groups. Chapter 11. Special Behaviour

The species interaction within a group is distinguished by a certain animal hierarchy. Moreover, such interactions are common both among mixed herds where strong adult bulls prevail and among solely male or female groups. The leader of multi-age female groups with calves is chosen by the size and age. As a rule, it is a large and experienced cow of old age. Sometimes, cows fi ght for the leadership in a group. Confl icts are often started by old females that cannot with- hold leadership and try to oust certain species from their group. Such confl icts provoke and disturb the entire group and even may lead to further confl icts. Such fi ghts might result in injuries and even fatalities. The leading female mostly gov- erns the life of a group defi ning the time for migration, choosing a grazing loca- tion and places for rest, etc. Hierarchy and Interrelations within a Group. The observers found a mixed group feeding on the slope of the Byky-ayagy isolated terrain in April 1970. At 10 o’clock, small groups of two to three species broke the main herd, walked a small distance away but stayed close enough to the main herd. Some animals stopped feeding and just stayed where they were waiting for the group to move. The herd moved after an adult bull started moving towards the fi r forest only; the herd followed it and came out of sight. Many group members motivate the leader to make a decision with their be- haviour. Sometimes, when two groups came together, or several animals joined a group, some animals, mostly bulls, had a rest or went to watering holes sepa- rately. Probably, joining a herd by a species does not mean its recognition by the other members. Although formally they are members of such group they still live separately, sometimes keeping within the group and sometimes not. The leader’s infl uence is more distinctive in stable groups of cows and calves. When such groups come together, and a newly appeared cow claims the lead- ership the leaders perform the ceremony of acquaintance, including thorough visual examination, sniffi ng, head shaking, sideways walk around the opponent, and demonstrating threatening positions. Often, it is enough to defi ne the new leader of such unifi ed group. However, when the power, age and size of oppo- nents is similar such ceremony of acquaintance often ends up in a fi ght, upon the results of which the defended cow might be ousted from the herd or take a lower position in the society. Such merger often does not lead to any distinctive Chapter 11. Special Behaviour 143 confl icts or fi ghts for the leadership. For example, in autumn 2001, a mixed group of cows and youths (all in all, fi ve species) moving from the Bugoy-chat isolated terrain met four wisents feeding not far from the path in the spruces of the Sofi yskiy Mountain Range. After the fi rst group came up the slope and got seen by the other group, the wisents stopped and looked at the feeding animals for a while. The second group took a notice of the fi rst one and kept feeding, only an adult bull kept watching the new group approaching. The “guests” also started feeding and moving towards the coniferous forest in the upper part of the slope. The bull broke from its group, came to the new animals and started sniff- ing some females; the groups started going closer to each other. When they came close enough the animals stopped again and observed each other for a while, then a female from the second group slowly started moving towards the conifer- ous forest and others followed it almost in a single group. There were no signs of antagonism seen among the animals. It is worth mentioning that the bulls and cows do not compete for the leader- ship; moreover, adult bulls often act as “peacekeepers”: with a loud snort and raised tail they interpose between females and take them apart (Danilkin, 2005).

Figure 66. Close Interaction of the Species Plays a Significant Role in the Life of the Wisents. Photo by U. Semenov. 144 U.A. Semenov The Wisents of Karachay-Cherkessia However, relations of bulls and cows change in the periods of pregnancy and lactation. In June 1992, some employees of the Nature Reserve saw a cow get aggressive when a young bull approached its calf; it immediately made the young bull go aside. The cows got more aggressive before and after the partu- rition due to the need of solitude. When they rejoined the herd with grown-up calves they became calmer as their offspring immediately got under protection of other wisents. Competition and Interrelations with Predators. The wisents have special re- lations with bears. Within the fi rst years upon the wisent release in Arkhyz, the observers several times saw bears approaching the wisent herd in spring and sum- mer; they also saw some unfortunate attempts of the bears attacking the wisents. Afterwards, the wisents were not afraid of those predators and let them come closer without running or showing any signs of disturbance. As a rule, the bears did not attempt to attack the wisents; they just came closer to a herd, thoroughly inspected it (probably, assessing the state of the animals and trying to fi nd some ill and weak species) and walked by, while some bears even stayed and fed fresh grass next to the feeding wisents. A. Batchaev, an employee of the Nature Reserve, several times saw a bear feeding at the distance of 40–50 m from the wisent herd in April and May 1973. The animals were very calm and did not pay any attention to each other. However, the appearance of the bears in the areas where young grass grows in early spring leads to the competition among them and the wisents. As the wisents’ diet is similar to that of the deer and roe deer, and less similar to that of the wild boar (the wisents graze on 90.8%, red deer on 94.6%, roe deer on 91.3%, and wild boar on 48.7% of all the plants in the diet of the ungulates; moreover, the wisents feed on 5.1% of the forage eaten only by them) (Zablotskaya, 1957; Alexandrov, 1958; Durov, 1983; Dyrenkov et al., 1990; Kazmin et al., 1992). The feeding competition of the species strengthens under unfavourable winter condi- tions (Shostak, 1973). Some scientists believe that despite so many diet similarities there is no real competition between them as such species prefer different plants (Zablotskaya, 1957; Alexandrov, 1968). The observers of the Caucasian Nature Reserve believe (Nemtsev et al., 2003) there is no real competition between the wisent and the deer. However, in the Bialowieza Forest, the deer reduces the envi- ronmental niche of the wisent in winter (Shostak, 1973). According to our researches in Arkhyz, the competition of the ungulates is more severe in certain stations. Sometimes, the wisents and deer fed simultane- ously at the same forest glades. The island effect of some feeding stations (salt springs, sun heated slopes with earlier grass appearance, fruit trees, avalanche cones, etc.) of the mountain environmental systems force the animals to come thereto, thus, resulting in enhanced competition. Social Behaviour as a Form of the Landscape Adaptation. The behaviour of the wisents changed upon their exploration of the new territories and biotopes Chapter 11. Special Behaviour 145 of the Kizgych Valley. Besides, in 1977–1978, about 30% of the animals used to leave the Nature Reserve and to stay in the isolated terrains of Psyzh, Sofi a, Rapachay, etc. for a long while. In summer 1978, the entire herd left the Kizgych area in several groups for a short while and even fed with domestic cattle on the summer grazing lands. It was mostly seen close to the Sofi yskoe Sedlo Mountain Pass. Moreover, as it had been previously mentioned, the wisents never stayed overnight in the alpine zone. Instead, they left for the forest and next morning came back to the upper border of the alpine meadows for feeding. They stayed there until the evening, preferring windswept areas in order to keep away from gnats and fl ies. They came down the slope in the evening. The altitude range of their migrations and the duration of their stay in the open areas within the warm season mostly depended on the weather conditions. On sunny days, they came higher and stayed there longer than on dull days. In case of nasty weather they browsed and rested next to the upper border of the forest and subalpine areas. It is hard to say that the animals quickly explored the entire Kizgych area, though they came to the avalanche cones already within the third year upon their release. At fi rst they started coming there in summer, then used them as feeding stations in early spring grazing on young branches and bark of snow bent trees and shrubs. Such areas are mostly visited by the bulls. In a while, the wisents changed their migration pattern within the chine (Tara- sov, 1980). Previously, the wisents moved all other the valley from the river mouth up to the midstream within the entire winter, but the snowfall of March 1973 limited their migration routes, so they moved within the fl oodplain up to the glade of Perviy Post. There was a sharp border between the moderate (40–50 cm) and deep (50–80 cm) snow at that place. The daily migration distance got four to fi ve times shorter already in the middle of December as at some points the snow depth reached 1.4 m. When the snow cover was thin they walked 6–7 km, but in deep snow the length of their daily migration routes did not exceed 1–1.3 km. It is worth underlining that predators immediately tried to use severe life condi- tions of the ungulates. Wolves tried twice to attack two wisents that kept apart from the groups in December of that year. The observers failed to determine the wisents’ gender and age but approximately they were two bulls — an adult and a young one. The animals skilfully defended themselves by standing close to each other, thus, none of them got hurt. Behaviour during the Rutting Season. The wisent interrelations changes as the rutting season begins. At such time, there is a sharp aggravation among the bulls that get aggressive and agitated, often fi ght and even chase their opponents in short-distance runs. Although confl icts of males and females over the social leadership happen not only within the rutting season, they become much more frequent at such time. The species confrontations start with mutual examination, threatening at a distance (threatening positions and head shaking) and then ap- 146 U.A. Semenov The Wisents of Karachay-Cherkessia proximation. Sometimes a bull takes it out on the shrubs and young trees furi- ously rubbing against them. If the power difference is signifi cant the wisents use only distant threatening tactics, as a result of which a weaker bull leaves. A confl ict of two equally strong bulls often leads to a severe fi ght. The animals butt heads and hit each other with their legs trying to reach the opponent sideways. Sometimes the blows are so hard that an opponent falls on the ground, and such injuries might result in fatalities. Thus, in August 2000, a fi ght of two bulls of the similar age and weight resulted in the death of one of them in the Wisent Nurs- ery of the Okskiy Nature Reserve (Tsibizova, 2004). Sometimes several young species ousted by an adult bull come together and attack it bringing it to death (Danilkin, 2005). Behaviour Change Dynamics. The wisent dynamics gradually changed along with appearance of the animals born in Arkhyz. The calves born there were more cautious than their parents and, in a while, more seldom showed the behaviour patterns of the animals released in that area. For example, the released species were not afraid of the noise made by the vehicles and tractors that brought hay. They did not run away hearing a human voice at the avalanche cones or a power saw noise. They stayed calm when a bulldozer smoothed paths 10–15 m away from them. Nearly complete lack of human persecution, as well as “conven- tional acquaintance” of the released animals with people resulted in high level of adaptation to the manmade conditions. The animals easily came to the village borders and roadsides, and stayed in the areas of holiday camps, holiday houses and even commercial dairy farms for a long while. Such occurrences were regis- tered even in 1987. Once, two wisents lived next to a holiday camp for the entire winter. Some species even came to the village at night and ate the hay in private households. Moreover, two sexually mature bulls several times came to a cattle farm of the Shorovo isolated terrain and mated with domestic cows. It is known that three hybrid calves were born as a result of such breeding (Bobyr, 1992). Some local people insisted that young cows mated with wisents could not fawn without signifi cant human assistance, though some adult cows did it without such assistance. However, such data require additional clarifi cation. The behaviour of the calves of the current year looked different in comparison to their “manmade” parents. They were very cautious; elder calves ran away from the herd upon approximation of people and stopped, then ran away a bit farther and stopped again as if trying to make the others follow them. Some- times, the herd saw the disturbance of the calves, so the cows with calves left the groups in order to keep away from vehicles and people. Sometimes such groups were followed by a bull. We studied the Teberda Reserve Nature Records for 1968–1978 and found out that the cows nursed the calves born a year before un- til late June. It was found out that the calves were nursing for about 10–12 min- utes in the beginning of the lactation period of the previous year, while at the age Chapter 11. Special Behaviour 147 of 11–12 months they were nursing for three to four minutes in the end of such period. Small groups of two to four species come together building larger herds of seven to over 16 species in hard winter periods, especially when there is a lot of snow. It facilitates their walk through the deep snow as the strongest animals lead the way. For example, such groups were formed in the winter of 1975/1976 when there was a lot of snow. Building larger herds is signifi cant for the popu- lation’s survival in the mountains. Cows and youths can follow stronger bulls (sometimes, adult cows) and do not lose too much strength on searching for food. The animals survived that winter well; by the spring the cows looked much better-fed than the bulls. It was found out that the cows had been searching for food more actively than the bulls. The observers pointed out that those were the cows that broke the fence around the hay stacks forcing the way with their chests or prying it up, thus reaching for the food earlier than the bulls. If the animals failed to break the fence they tried to jump it over, sometimes overcoming ob- stacles at the height of up to 170 cm. At such time, the wisents become less cau- tious and often access the hay stacks located next to the houses or roads. In spring 1978, the release of a new group of seven animals (four bulls and three cows) into the Kizgych Valley changed the social structure of the Arkhyz wisent population. As a result of their appearance a new herd was formed that did not fall apart for a long while. Only one young bull (a newly released one) was driven out by a “local” adult bull and left for the upper part of the neighbouring Shorovo Chine where it stayed until October. The observers saw it sometimes grazing together with the domestic cattle. After the cattle left the wisent came to the mouth of the Psyzh River for wintering where it stayed at logging plots of the Arkhyz isolated terrain until mid-January 1979. It is not clear what happened to it later, probably it joined the herd. In three years, one more bull left the nature reserve for the border of the Psyzh isolated terrain, from where it moved towards the village of Pkhiya. Shepherds several times saw a wisent grazing next to the path leading to the mountain pass in late July. They say that in autumn, in the third decade of September, that wisent was seen in the area of the Temir-kulak Mountain Pass. Afterwards, no traces of it were found. It was not clear whether that wisent was the one ousted from the herd or another bull, or what happened to it afterwards. Interrelation with People. Depending on their condition in different seasons, the wisents show different behaviour patterns towards people. They get more aggressive and intolerant to people within the rutting season and close to the offspring (Tarasov, 1977). In September 1973, a bull attacked an observer who incautiously came too close to a herd with a calf of the current year (the wisents were not afraid of people at that time and let them come quite close, so the wi- sents got anxious when people appeared 20–30 m from them). The wisent did 148 U.A. Semenov The Wisents of Karachay-Cherkessia not chase the observer, came back to the group and kept watching the observer. It got quiet and started grazing in a while. As a rule, the animals leave their place when they feel people approaching and try not to let them come too close, but sometimes curiosity wins over the caution, and observers manage to come quite close to certain species. In the course of the area monitoring in May 2012, we saw a solitary adult bull in a thick deciduous forest of the fl oodplain terrace of the left bank of the Kizgych River at the distance of approximately 85–90 m at approximately 06:00 p.m. The observers rode horses up the path, and the wi- sent looked at them without trying to escape. However, 15–20 seconds after the observers stopped it went 10–12 m away and stopped. When the observers dis- mounted and tried to come closer the wisent rushed towards the forest breaking the tree branches on its way. We saw another group of wisents on a wide glade of the southern slope of the Sofi yskoe Sedlo Mountain Range at the altitude of 2,000 m above sea level at about 07:00 p.m. of the same day. A group of four species (adult bulls aged 9 to 12 and 9 to 10 years, respectively, a cow aged seven to eight years and a young bull aged two years) was grazing there. As the animals saw the people they got cautious but did not try to escape and stayed where they were. The observers stopped at the distance of 650–700 m from the animals and looked at them without dismounting. The cows started grazing in a while but the bulls kept looking at the observers. The wisents regrouped at the sight of a dog. The bulls moved ahead and stopped at the distance of 10–12 m from each other. The dog approached the fi rst bull and started circling around it with loud barking. The second bull immediately approached the cows, thus, forming a circle, in which the animals stood back to back. The second bull kept staying where it was without trying to leave or join the group. The wisents were observing the dog but in a while the second bull (a younger one) left the cows and started chasing the dog at fi rst in short and then in large headlong jerks of 5–10 m. After the dog left the wisents started up the slope grazing on their way. The next day, on May 5 at 08:00 p.m., an adult bull previously seen on the forest glade was found at the altitude of 2,200 m. It kept grazing even upon tak- ing notice of an observer at the distance of about 450–500 m, from time to time it raised the head and looked at the surroundings and the observer. I dismounted and slowly approached the wisent moving towards it while it was grazing, and stopping when it raised the head and looked in my direction. When the distance between me and the wisent decreased to about 100 m, I started moving more slowly but without breaks. Thus, I shortened the distance down to 60–65 m be- fore the wisent started escaping. The wisent got attracted by the photofl ash, so it stopped and kept looking within a long period of time. It allowed me to reach the distance of 20–22 m before the wisent started escaping but time and time again it stopped seeing the fl ash. In a while, it got bored and quickly went towards the forest. Chapter 11. Special Behaviour 149 Therefore, we assume that the wisents feel more confi dent in the open moun- tain areas as they can monitor the surroundings in case of appearance of people or predators. Besides, sometimes they stay still just looking at people but then quickly leave and sometimes sneak from their place upon sniffi ng people, and only the sound of breaking branches indicates the direction of their run (Se- menov, 2012). Despite the slow naturalization of the new Arkhyz wisent group, some species still remembered their acquaintance with people, which used to play a signifi - cant part in their life within a long period of time. The wisents were not afraid of people even 15 years after the release. They easily came to the deforestation areas, approached people, and fed on the bark and needles of the woods until the middle of the 1980s. Chapter . 12. Key Extinction Reasons

The adult healthy wisent is a large and strong animal rarely attacked by preda- tors. Powerful horns, heavy weight, strong legs, and quick reaction allow bulls and cows to repel attacks even of a group of predators. Still, young, ill and weak animals often fall victims to wolves, bears and lynxes, while thick snow and ice crust only increase the possibility of their successful hunting. After the wisents’ release in the Kizgych Valley in 1969, A. Batchaev, a wood- ward, saw a bear attacking wisents, and later on a wolf coming close to a group of animals and then leaving. It is also known that a group of wolves once chased several wisents. In April 1987, after a cold and snowy winter, remains of a wi- sent eaten by a bear were found in the fl oodplain (Bobyr, 1992); however, the observers failed to defi ne whether that bear had attacked a weak animal or just scavenged. Besides, no wisents were killed by bears or lynxes in the area of the Caucasian Nature Reserve. Still, E.V. Pfi zenmayer (Pfi zenmayer, 1929, cit- ed by Nemtsev et al., 2003) wrote that wisent remains had been several times found in the stomachs of bears. According to N. Kutepov (1911), a bear killed fi ve wisents within the summer of 1846 in the Bialowieza Forest. Three bears at- tacked an adult bull in the Nalchik Hunting Reserve (Kalugin, 1968). According to some researches (Bashkirov, 1939; Rashkevich, 1973; Bobyr, 1980), lynxes and leopards also hunt calves, but it happens really seldom. A. Kudaktin (1982) believes that the presence of wisent fur in the excrements of a wolf in the Cau- casian Nature Reserve proves that predators scavenge. According to the observ- ers, a bear attacked a young cow aged three to four years in the Tseyskiy Nature Reserve (North Ossetia) (Bobyr, 1980). In the course of the examination and recording in the Kizgych Valley in No- vember 2000, we found traces of a multi-age group of fi ve to six wisents in the midstream heading to the upper part of the chine. Those traces were followed by the traces of a group of three to four wolves. In some distance, the observers found some fragments of a wisent coat and bones not far from the salt licks. It was impossible to defi ne the gender and age of the wisent but it was supposed that it was young. Chapter 12. Key Extinction Reasons 151 In April 2003, some fragments of carcass and coat of an adult wisent fallen in an avalanche were found. The wisent was partially dug by a bear whose traces and excrements were found not far from the place where the wisent had died. The predator had been coming to that place and scavenging for a few days by the time the observers found the remains. It was impossible to defi ne the gender of that wisent. Some fragments of a wisent coat and fur were found in the avalanche cone in early April of the next year. A bear dug a hole in the dense avalanche snow and found the wisent at the depth of 4 m. It was supposed that several wisents had died in that avalanche but, unfortunately, it was impossible to repeat investiga- tion of that area and defi ne the gender and age of the fallen animals. In late April 2005, a dead adult bull was found in the upper reaches of the Kiz- gych River. Its remains partly eaten by a bear were found within the avalanche cone. Wolves successfully attacked a wisent at the salt lick only once within the entire observation period. At that time, the observers found some fragments of a wisent coat. Still, it was not clear whether the animal had been ill or weak. The wisents of the Caucasian Nature Reserve also seldom suffer from the wolf at- tacks; the wolves successfully hunted only two calves of the current year within 25 years (Kalugin, 1968). Therefore, 33 wisents died in the area of the Kizgych Chine of the Teberda Nature Reserve (Table 13) between 1968 and 2012. The remains of 30 of them were found and investigated. Two animals had injuries and afterwards were seen neither within any group nor separately. They were deemed dead outside the Nature Reserve. Most of the Arkhyz wisents died from emaciation and weakening due to snowy winters (34.4%). The area limitations and impossibility to migrate to less snowy areas make this factor the key reason for population development restraint. The Arkhyz wisents also suffer from different diseases. Domestic cattle dis- eases brought to the mountains in the past became one of the reasons for the Caucasian wisent extinction; such diseases remain an acute problem in the present. It is known that at the end of the 19th century the animals suffered from the hoof-and-mouth disease, and later from the anthrax (Dinnik, 1910; Bash- kirov, 1939). All in all, the wisents are sensitive to the domestic cattle diseases. Therefore, the wisents more often suffer and die of the hoof-and-mouth dis- ease, pasteurellosis, anthrax, brucellosis, tuberculosis, blackleg, paratyphoid fever, pneumonia, tetanus, rabies, actinomycosis, etc. (Vrublevskiy, 1908; Dinnik, 1909; Severtsov, 1926; Bashkirov, 1939; Kalugin, 1968) in many ar- eas where the wisents live close to the domestic cattle (Bialowieza Forest, North and North-Western Caucasus, and Askania-Nova) or when wisents live 152 U.A. Semenov The Wisents of Karachay-Cherkessia Table 13 Reasons for the Wisent Extinction in the Teberda Nature Reserve and Number of Dead Animals (According to the Teberda Nature Reserve Records for 1968–2010)

Gender and Years Age of Dead Cause of Death Number Animals Fracture of a fore and a hind limbs in the course of Adult cow 1969 passing an area of large rock debris 2 Adult cow Fallen down a snow cornice on the cliffs

Died in the avalanche in the Bugoy-chat isolated terrain Adult cow at the upper forest border

1975 Died of a disease in the fi rst decade of July. The animal 2 Adult bull was not moulted; it was hidebound and had had a severe cough within its last days.

On July 23, remains of an adult bull were found in the fl oodplain of the Kizgych River. The wisent was very hidebound, half its teeth were lost, another half 1976 Adult bull severely worn, bare and mercurial. The autopsy showed 1 that it had died of a malignant tumour of the right cerebral hemisphere, hemorrhagic stomach and bowel infl ammation, and cachexia.

Died in December despite the treatment by the Animal Health Service and Forest Service. The autopsy carried out by a veterinarian did not help to defi ne the cause 1977 Adult bull of death. The wisent had been ill within half a year, 1 had moved too little and had not stood up before the death due to the hind limb paralysis. It had been very hidebound and suffered from cough.

Six bulls left the Nature Reserve in March. Among them there was a bull with an injured horn and a tumour in the Adult bull pelvic area. Though its remains were not found it was 1 not seen within that group. It was supposed that it had 1982 died outside the Nature Reserve. It used to be a member of a group and limped hard. Though its remains were not found it was not seen Adult bull 1 within that group. It was supposed that it had died outside the Nature Reserve due to a hoof trauma.

1985 Adult cow It suffered from emaciation and severe cough. 1 Chapter 12. Key Extinction Reasons 153 Table 13. (continuation) Adult species (gender not Emaciation due to a cold and snowy winter. 1987 defi ned) 3 Young cow Emaciation due to a cold and snowy winter. Adult cow Emaciation due to a cold and snowy winter. 1988 Adult bull Disease (more exactly not defi ned) 1 Disease (weakening due to the old age of more than 1989 Old bull 1 20 years, severe limp) Species (age 1990 and gender not Cause of death not defi ned 1 defi ned) Wisent remains were found between January and June, Bulls and cows 1992 the animals were very hidebound due to a cold and 8 of different age snowy winter. Disease (weakening due to the old age of more than 20 1993 Old bull 1 years) Adult species 1996 (gender not Not defi ned 1 defi ned)

In April, an adult limp cow was seen within a group. 1998 Adult cow It looked emaciated and kept separately from the herd. 1 Afterwards, it was not seen, thus, it was deemed dead.

Adult species (gender not Died in the avalanche (there was found a skull) 1 defi ned) 1999 Adult species (gender not Died in the avalanche (there was found a skull) 1 defi ned) Species (age Hunted by predators (there were found coat fragments 2000 and gender not 1 not far from the salt lick) defi ned) Adult species 2003 (gender not Died in the avalanche 1 defi ned) Species (age 2004 and gender not Died in the avalanche 1 defi ned Adult 2005 species(gender Died in the avalanche 1 not defi ned) 2009 Adult cow Died in the course of parturition 1 154 U.A. Semenov The Wisents of Karachay-Cherkessia in yards (which results in helminth contamination due to limited grazing areas and irregular fence replacement (Nazarov, Kiseleva, 1980). Many researchers point out that epizootics of the hoof-and-mouth disease, anthrax and pasteu- rellosis were most destructive for the wisents. According to Kalugin (1968), there were two pasteurellosis outbreaks in the Caucasian Nature Reserve due to the release of a wisent brought from the Moscow Zoo in 1941 and proximity of the domestic cattle grazing close to the Nature Reserve in 1955. Twenty- fi ve percent of all the deaths of the Arkhyz wisents for 44 years were caused by the diseases, while in the Caucasian Nature Reserve it amounted only to 18.2% among the calves of the current year for 25 years (Kalugin, 1968). However, it is worth mentioning that some diseases and deaths were caused by weakening due to snowy winters or injuries. Two dead animals that were found by observ- ers looked ill but their further investigation showed that they were of old age (over 20 years), so that it would be wrong to say that they died of a disease. Two animals underwent autopsy though it did not show any signs of the above epizootics. It is supposed that by the time of wisent release in Arkhyz in 1968 the domestic cattle had been well looked after, and the possibility of epizootics was minimized. Still, some domestic animals got ill. We believe that not all of the diseases were identifi ed as not all the dead animals underwent autopsy or were suffi ciently investigated. If the wisents and domestic cattle used the same pasture lands along the Nature Reserve borders within 44 years it is next to impossible that no diseases were caught. Moreover, the wisents of the Nature Reserves regularly died of diseases. However, the data available do not allow for fi nding exact interrelations. In terms of the wisent death from non-contagious diseases, the wisents suf- fered from parasite worms signifi cantly less than the domestic cattle. Moreover, the helminth fauna of the wisents does not include echinococcus and other ces- todes that are common for other ruminants (Pupkov, 1966; Rukhlyadev, 1967). Many Arkhyz wisents died in the avalanches (18.6%). The actual number of the wisents died in the avalanches might be even more signifi cant as they use the avalanche cones as feeding stations (even in winter), and it is hard to fi nd all the dead animals in such areas. As said by Vladimir Taloyan, who worked for the Caucasian Nature Reserve for a long while, he found some wisent coat and limb fragments in the avalanche cones several times. Injuries that sometimes lead to fatalities constitute another reason for the wi- sent extinction in the mountain environmental systems. All in all, about 12.5% of the dead wisents fell due to injuries. It is worth underlining that that indicator of the calves of the current year of the Caucasian Nature Reserve that has simi- lar environmental conditions amounted to 12.2% (Kalugin, 1968). The wisents often injure their limbs in the course of passing through the large scree debris, coming up and down scarp slopes. Two wisents died within the fi rst year after the Chapter 12. Key Extinction Reasons 155 release as those lowland animals had not been used to the mountain landscapes (snow cornices, cracks, avalanches, etc.). Afterwards, the observers also found some injured wisents (limp, injured horns and hooves, etc.) but not so many. It is possible to admit that injuries caused an insignifi cant number of deaths and did not infl uence the population survival, though every death counts for the entire micropopulation. Although the wisents sometimes cross the river within the summer river fl ood in the Kizgych Valley caused by the snow melting, no wisents died in the course of the river crossing, while the number of deaths of the wisents of the Caucasian Nature Reserve amounts to 12.2%. Moreover, no wisents died in the course of intergroup fi ghts although some species had broken horns and had other injuries. Still, the competition among the bulls was severe as young bulls sometimes were ousted to the outskirts within the rutting season. Some of them left the groups and did not return gradually going far away from the place of their initial release. It is not clear what happened to those animals afterwards. Adult bulls often die during the mating season when the competition strength- ens. For example, two bulls of the Nalchik Hunting Reserve got severely injured in the course of a fi ght, one of them died (Tsaryapin, Vysokov, 1963). The num- ber of the wisents dying in fi ghts reaches 10.9% in the Caucasian Nature Reserve (Kalugin, 1968), and 12.5% in the Bialowieza Forest (Wroblewski, 1927). Be- sides, in the course of fi ghts, the bulls injure themselves against the rocks or by falling. Some cows die in the course of parturition or deliver dead calves (Koroch- kina, 1971; Vasilyuk, 1973; Kalugin, 1973; Kiseleva, 1974; Bunevich, Koch- ko, 1988). In 2009, a wisent cow that had died in the course of parturition was found. Its carcass with an unborn calf was partially eaten by a bear. It is supposed that the cow could not deliver a calf and died; afterwards, it was found and eaten by predators. Young wisents and calves often die at the feed- ing places where stronger species trample them to death in front of the hay stacks. Some wisents die from intoxication. Some researchers insist that some animals died from tympanitis due to the over-consumption of oat and beet, as well as fatal poisoning from eating the European white helleborine and ranun- culus acris (Kalugin, 1968). The mountain poaching is an acute problem. Although until the end of the 1990s the Arkhyz wisent population had not suffered from severe illegal hunt- ing pressure the way the Caucasian Nature Reserve did (Nemtsev et al., 2003; Danilkin, 2005), some wisents might had been hunted. As said by G.Ya. Bob- yr, an employee of the Teberda Nature Reserve, he accidently found suspended wisent carcass and some carcass and coat fragments on the ground next to one of the camping sites in 1988. The wisent came close to the camp and was shot down (probably, it was caught and then slaughtered as at that time the wisents 156 U.A. Semenov The Wisents of Karachay-Cherkessia were not afraid of people, and it was easy to allure one and throw a loop of rope around its horns). Still, single cases could not signifi cantly infl uence the population dynamics. The population state dramatically changed upon the en- try of soldiers to the upper part of the Kizgych Valley. In 2000, the qualitative reporting showed the catastrophic consequences of their stay in that area for several years. Conclusion

The fi rst observations we conducted in 2000 showed that the wisents did not keep to the headstreams of the Kizgych River. All the cases of seeing them were registered either in the middle stream of the river or in the lower territories. The reasons for that phenomenon were unclear at that time because according to the data of the Teberda Reserve Nature Records for the previous years, the Bugoy- chat isolated terrain and the Sofyiskoe Sedlo Mountain Range spurs used to be one of the permanent locations of the animals. Our observations did not confi rm that. Moreover, other animals were also seen there infrequently. The wisent population check carried out that year revealed a discrepancy be- tween the offi cial data and the actual situation. There was no massive poaching by the local people. However, one cannot totally rule out the possibility of some species crossing the reserve borders and being shot down though it would have been impossible to conceal the killing of 26 species (according to the check in 1999 — 46 animals; according to the check in 2000 — 20 animals). Various reasons for such an abrupt drop of the population were considered: a group of animals being killed by an avalanche, animals leaving the reserve, short-counting of animal groups and all the above factors collectively. These versions should have been confi rmed sooner or later by the animal corpses found or by reports of shepherds, hunters, mushroom hunters or tourists about see- ing wisents in other territories the way it used to be during the previous years, but none of that happened. Moreover, the wisents left the reserve less and less frequently from year to year. And only a few years later, it was revealed in the course of unoffi cial conversations with some military people and their local guides that there had been poaching cases in the late 90s. The near-border territory regime provided for a convenient mechanism of re- stricting the access to the Kizgych Valley, which helped to prevent undesirable people from coming there, including the reserve staff. The author himself en- tered into numerous confl icts with the military offi cers because of their multiple violations of the reserve regime and attempts to keep him away from the upper part of the chine that he was visiting for the purposes of performing his duties. That relatively short one or two year-long impact on the Arkhyz population that had been restituted for over 40 years had such catastrophic consequences that the population did not manage to recover on its own even after the pres- sure had been relieved. Furthermore, the presence of the contract soldiers in the Kizgych Valley caused the dispersion of the animals and ousted them from their 158 U.A. Semenov The Wisents of Karachay-Cherkessia comfortable habitual places to the other less favourable environment where the probability of their death due to the natural factors that time (thick snow and avalanches) increased drastically. The population dynamics, birth rate and habitat structure of the animals’ spread during all the seasons were to be analyzed for the period before the military came there in order to evaluate the stability of the population development and to sum up the results of the species reacclimatization in Karachay-Cherkessia, because there had been no active human pressure on the animals, and their population had been developing on its own until that moment. As described in Chapter 3, the abrupt drop of the wisent population was regis- tered in 2000. We suppose that the tendency had started before because the mili- tary had been staying in the upper part of the Kizgych Chine for a few years by then, while the count of the animals had not dropped because of the population overestimation by mistake. In those years, they roughly estimated the number of wisents outside the reserve borders and summed it up in follow-up of the winter check based on the traces and visual contacts. That resulted in the double count- ing of some species because the animals could easily come back to the check territory within several hours (the checks of the wisents in Arkhyz were usually carried out for two to three days by way of visual examination of the river fl ood- plain and of some accessible isolated terrains) or, on the contrary, they could leave it creating an illusion of more groups existing. Such mistakes were inevi- table taking into account that the total number of the animals was calculated by the method of summing up the animals visually registered in some places and the traces found in other places, while the number of animals within the groups often changed in the course of migrations, which complicated their identifi cation. The quality of the checks carried out in some years was also doubtful. The problem of counting the wisents in the mountains is still open. The cur- rently used practice of group registration in Arkhyz in winter has a number of drawbacks, such as highly probable double counting, work complexity during the winter period, and some animals being outside the reserve borders. The method of counting the wisents in the mating aggregation places used in the Caucasian Reserve is complicated due to the short term of the aggregation and the continuous migrations of the animals. During that period, adult bulls join the herd ousting the younger bulls, which leads to constant changes in the quantita- tive composition of the groups. Nevertheless, the method, if applied properly, makes it possible to register 80–85% of all the animals within three to four days (Nemtsev, 1980). The method is convenient for and applicable to big popula- tions and wide territories. The error does not exceed 5–8 % for micropopulations consisting of 50–60 species. In case the number of animals decreases it is impor- tant to count every wisent, which requires a comprehensive method of counting the animals throughout the year, including in spring on the open glades that the Conclusion 159 wisents visit to graze on the new grass. The experience showed that modern electronic methods of registering the animals help to quite accurately identify them and count each species. Wild ungulates start actively visiting natural and artifi cial salt sources in spring which is the best moment for counting the wisents after winter by installing hunting cameras in such places. The same method can be used to count the animals on feeding sites during the winter period. But only combining all the above methods will help to minimize the error of the checks and to assess their population dynamics objectively. The study of the archives of the Teberda Reserve Nature Records showed that the peak of the wisent population growth started in 1978 after a new group was brought in and continued till the early 90s when the population degradation signs became more visible due to the increased inbreeding factor refl ected in the decrease of the animals’ sexual activity during the mating season and in the increased rate of farrow females reaching 92–94% in 1994–1996. The rate of the wisents’ migrations to neighbouring areas had grown consider- ably by that time. Although the tendency of the animals crossing the Kizgych forestry borders had started earlier, it became stronger in the course of time. As their population grew the animals had to look for new territories outside the re- serve more frequently. The situation was complicated as the grass on almost all the accessible meadows and glades outside the reserve was cut (around 150 ha), which resulted in destruction of the most important winter feeding stations every year. At the beginning, when the number of the wisents was low, that impact was not noticeable but then the defi ciency of winter pastures became a very acute problem. The practice of large-scale hay cutting in the Kizgych Chine not only damaged the winter feeding base of the animals (in summer, the animals often left for the subalpine and alpine zones where they were never out of food) but also created a long-lasting wildlife disturbance. Hay stocking works were car- ried out there using specialty vehicles for one-and-a-half to two months, includ- ing hay cutting and transportation. The complex combination and interrelation of the ecological and anthropogenic factors led to reduction of the reproductive rate, disruption of the gender and age structures of the herds, and growth of the animals’ migration activity. Sustainable development of the population could be achieved by way of bringing in new wisents and taking comprehensive measures to ensure the ani- mals’ spread, including through expansion of the territory of the Teberda Re- serve Arkhyz area. However, the changes in the situation inside the country and the diffi cult economic and political circumstances in the Caucasian Region prevented those plans from being implemented. The combination of historical circumstances that left the problem of the species preservation unattended for many years and got even more complicated by the poaching, drove the Arkhyz population of wisents to the verge of total extinction… 160 U.A. Semenov The Wisents of Karachay-Cherkessia It is noteworthy that problems of restitution of the Caucasian micropopula- tions, including the Arkhyz one were touched upon during the discussion of the strategy of the wisent preservation in Russia in 2012. Some scientists expressed their doubts whether it would be reasonable to bring in new animals to the Kiz- gych Valley because it had no prospects for big herd formation required for their self-sustaining development. A rightful question arose: was there any sense in bringing in new wisents when it was clear from the very beginning that it would be impossible to form a big population there? There is certainly no sense in it if the wisent preservation in Karachay-Cher- kessia is considered to be a one-off release of the animals only. The population will be doomed to gradual degradation and extinction. Unfortunately, non-inter- ference with the course of natural processes alone, even in specially protected natural territories, cannot solve the problems of the biological diversity preser- vation today unless the areas are large enough. Many species cannot exist with sustainability within the borders of small reserves or national parks. That is espe- cially relevant for large mountain animals the life cycles of which are related to

Figure 67. A Group of Wisents Consisting of the Animals Newly Brought in 2012 Walking to the Feeding Sites. Photo by R. Tekeev. Conclusion 161 seasonal and daily migrations off the controlled territories when winter feeding stations are defi cient. The Arkhyz area of the Teberda Reserve being an isle of relative prosperity cannot alone ensure the existence of a big population there, which actually should not be targeted at all in such conditions. For example, the analysis of the present day ecological conditions in Belarus showed that the most suitable mode of preserving the European bison is the metapopulation model stipulating establishment of seven to nine centers in different ecological conditions and development of mixed-quality (mixed-type) subpopulations. It is pointed out that in case a population of at least 50 species is kept in the wild, wisents tend to form typical gender-age and demographic structures ensuring the reproduction stability (Kozlo, 2004). Thus, forming small populations, including by way of bringing in new ani- mals repetitively and exchanging species with other small micropopulations is an important component of the general strategy of the species preservation in Russia. The Arkhyz population containing at least 50–55 species is quite capable of sustainable existence within such territories provided the right strategy of the population management is developed. That means that the wisent preservation in the Teberda Reserve must not be limited to one-off campaigns of bringing in wisents. There must be a comprehensive long-term program implemented step- by-step. The program must include continuous monitoring with assessment of the population development stability and of the animals’ condition, and a drastic change of the territorial security regime eliminating all the consequences of the military presence in the chine, while the stationary reserve security stations must be replaced with mobile ones located in the forest where they would not interfere with the animals’ migrations. Hay cutting must be stopped within the entire territory of the Kizgych forestry. In case hay is needed for feeding the animals it must be cut outside the reserve or purchased. In order to prevent in- breeding, the practice of regular bringing in a few animals of different gender and age groups at a fi ve to six year interval must be introduced to keep up the total population of at least 50 wisents. The above number of wisents can com- fortably dwell in the Kizgych and the adjoining territories having prospects for population growth and further spread subject to implementation of the de- scribed program components. The released animals have been developing new adaptation capabilities for decades handing down from generation to generation their ability to survive in the diffi cult and fast-changing environment of the Arkhyz mountain ecosystems. We believe that preserving small and medium sustainable populations of wisents is important not only because they are an integral part of the Caucasian fauna but also because they could become safety stocks for the species restitution in case of epizootic outbreaks or deaths of the animals in other climatic zones. 162 U.A. Semenov The Wisents of Karachay-Cherkessia The strategic and prospective way to maintain the biological diversity and economic development of the region is to establish a national park “Arkhyz” with the area of at least 300,000 ha sharing borders with the Kizgych forestry of the Teberda Reserve. That would provide for natural development of habitat corridors for the wisents’ migrations and their exploration of new territories. Creation of such a protected natural reserve of national importance will prevent defragmentation of the biologically valuable solid forests by the owners and can ensure implementation of long-term tourist and environmental protection programs, including preservation of the wisent and other rare species of the wild fl ora and fauna of Karachay-Cherkessia. References

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Moscow. P. 5–65 [in Russian]. Zablotskiy M.A. 1975. [Wisent Protection Principles, Approaches and Forms] // Ungulates of the USSR. Moscow: Nauka Publ. P. 242–243 [in Russian]. Contents Introduction ...... 5 Chapter 1. Evolution and Distribution of the Genus Bison ...... 8 Chapter 2. The Wisent in the Caucasus ...... 16 2.1. History of Studies of the Caucasian Fauna ...... 16 2.2. Studying of Wisents in the Caucasus ...... 18 2.3. Morphology and External Appearance of the Caucasian Wisent ...... 21 2.4. The Wisent in Karachay-Cherkessia ...... 24 Chapter 3. History of Release and the Population Dynamics of the Wisent in the Teberda Nature Reserve ...... 39 Chapter 4. Habitat Structure and Migration Dynamics ...... 63 Chapter 5. Biotopical Spread ...... 78 Chapter 6. Diurnal Activity ...... 96 Chapter 7. Diet ...... 102 Chapter 8. Social Structure Dynamics and Herds Behaviour...... 121 Chapter 9. The Wisent Reproduction and Calving Seasons ...... 132 Chapter 10. Moulting ...... 139 Chapter 11. Special Behaviour ...... 142 Chapter 12. Key Extinction Reasons ...... 150 Conclusion ...... 157 References ...... 163