The Leopard, Panthera pardus, (Carnivora: Felidae) and its resilience ...

The Leopard, Panthera pardus, (Carnivora: Felidae) and its resilience to human pressure in the Caucasus by Igor G. Khorozyan and Alexei V. Abramov Abstract. Widespread in the Caucasus until the mid-19th century, the Leopard, Panthera pardus, has become extinct in many areas of this region but is still able to survive in some others. We have compiled a database of 218 Leopard records dated 1861-2007 throughout the Caucasus (Russian North Caucasus, Georgia, Azerbaijan and Armenia). Of these, 77 records of killings, 4 records of photo-captures (10 pictures), and 8 records of scat origin proof by faecal bile acid thinlayer chromatography were used as the most reliable indicators of the Leopard’s presence. We discuss the history of the Leopard’s postglacial emergence in the Caucasus, its habitats, scales of eradication, trends in Leopard extermination measures and range shrinkage within the study period, in separate sections under each of the four countries of the Caucasus. All recent and current Leopard records in the Caucasus are confined to central and eastern parts of the North Caucasus, south-eastern and north-eastern Georgia, south-eastern and western parts of Azerbaijan, and south-western and southern Armenia. All these parts of the range, except for the Talysh Mts in Azerbaijan, must be connected with southern Armenia which, in its turn, is linked with the much larger population in north-western Iran. The Talysh Mts stand isolated from the Caucasus Mountains and are directly linked with Iran. Kurzfassung. Der Leopard, Panthera pardus, der im Kaukasus bis Mitte des 19. Jahrhunderts weitverbreitet war, ist in vielen Teilen dieser Region ausgestorben, kann aber in einigen Gebieten noch überleben. Wir konnten eine Datenbank mit 218 Leopardennachweisen aufbauen, die die Jahre 1861-2007 umfasst und sich auf den gesamten Kaukasus bezieht (Russischer NordKaukasus, Georgien, Aserbaidschan und Armenien). Von diesen Nachweisen beziehen sich 77 auf erlegte oder getötete Tiere, 4 auf Fotografien mit Fotofallen (10 Bilder) und 8 auf Kotproben, deren Authentizität mit Hilfe der Dünnschichtchromatographie bestätigt wurde. Wir diskutieren das Erscheinen des Leoparden im Kaukasus in der Nacheiszeit und die heutigen Habitatansprüche der Art. Für jedes der vier kaukasischen Länder stellen wir das Ausmaß der Verfolgung sowie die Trends der Gefährdungsursachen und die Verkleinerung des Verbreitungsgebietes dar. Alle kaukasischen Nachweise neueren Datums sind auf den zentralen und östlichen Nord-Kaukasus, Südost- und Nordost-Georgien, Südost- und West-Aserbaidschan sowie Südwest- und Süd-Armenien beschränkt. Alle diese Teilgebiete müssen, mit Ausnahme des Talysh-Gebirges in Aserbaidschan, mit den südarmenischen Vorkommen in Verbindung stehen. Diese wiederum stehen im Austausch mit der viel größeren Population im nordwestlichen Iran. Das Talysh-Gebirge ist vom übrigen Kaukasus isoliert, steht aber mit dem Iran direkt in Verbindung. Key words. Leopard, Caucasus, distribution, Georgia, Armenia, Azerbaijan.

Introduction The Leopard Panthera pardus (Linnaeus, 1758) has been living in the Caucasus since the Pleistocene and has evolved to co-exist with humans (VERESCHAGIN 1959). However, as human population growth became more and more evident in this region in the mid-1800s, this predator was increasingly perceived as an enemy and less space was left for its safe Zoology in the Middle East 41, 2007: 11–24. ISSN 0939-7140 © Kasparek Verlag, Heidelberg

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Zoology in the Middle East 41, 2007

existence. This was particularly true for the North Caucasus and Georgia which were the outposts of the Russian Empire in the Caucasus and where nature exploration and wildlife control endeavours were carried out most intensively. The eastern part of Azerbaijan has also been under substantial anthropogenic pressure due to the wealth of its natural oil reserves on the Apsheron Peninsula (GRID 2002). Armenia remained an undeveloped rural part of the Russian Empire and then of the USSR, and so scientific interest in the Leopard and other wildlife only began just before World War II. For this reason, Leopard records in the Caucasus cannot be used to estimate the numbers and their fluctuations over different periods of time since they reflect the levels of human activities and densities, which in turn affect the probabilities of man-Leopard clashes, and not the Leopard abundance per se. The total number of Leopards in the Caucasus is now estimated as maximum of 30 individuals (KHOROZYAN et al. 2005). Nevertheless, the species records provide a very informative tool to illustrate the general trends in population status and range structure in the region (Fig. 1). In this paper, we use the Leopard records to discuss the history of the postglacial emergence of the Leopard in the Caucasus, its habitats, the scale of eradication and trends in Leopard extermination measures, and range shrinkage from 1861 to 2007, in separate sections on the North Caucasus of Russia, Georgia, Azerbaijan and Armenia. We consider the Caucasus to be a geopolitical region which encompasses four independent countries (Armenia, Azerbaijan, Georgia and Russia) situated on the crossroads of Asia and Europe, between the Black and Caspian Seas. Its definition thus differs from the “Caucasus Biodiversity Hotspot” and “Caucasus Ecoregion” concepts which also include parts of north-eastern Turkey and north-western Iran (KHOROZYAN et al. 2005). The Leopard living in the Caucasus, northern Iran and Turkmenistan belongs to the Caucasian Leopard subspecies P. p. ciscaucasica (Satunin, 1914). The junior synonyms are Persian Leopard, P. p. saxicolor Pocock, 1927, and South Caucasian Leopard, P. p. transcaucasica Zukowsky, 1964 (KHOROZYAN et al. 2006).

Methods We have compiled a database of 218 Leopard records in the Caucasus from 50 sources, including the literature, personal communications, specimens in museums and in private collections, and our own studies. This includes 112 records from the North Caucasus of Russia (1875-2004), 45 from Azerbaijan (1866-2007), 41 from Armenia (1939-2007), and 20 from Georgia (1861-2005) which contain precise information about localities and time. General information such as “…the Leopard was particularly common before in the canyon of the Alazani River where it was often killed near Zakatali, Lagodekhi, Nukhi and Buma Canyon” (HEPTNER & SLUDSKY 1972, SLUDSKY 1973) was omitted. The geographic distribution of the records and the current range are illustrated in Fig. 2. The results of a statistical analysis of the records are given in Figs 3 and 4. Indirect evidence for the presence of the Leopard (tracks, roar, prey remains, scats, scrapes and dens) forms a significant portion of the predator records in the Caucasus (Fig. 3). However, in some cases they could belong to other animals, e.g. the Eurasian Lynx, Lynx lynx, or Grey Wolf, Canis lupus. Observations, especially at night, twilight or dawn, can also be mistaken and other large carnivores can be taken for Leopards. This is especially true for the Lynx, which attains a large body size in the Caucasus and is more common than the Leopard (HEPTNER & FORMOZOV 1941, own data). In order to estimate the dynamics and structure of the Leopard distribution in the Caucasus, we primarily used the data on killings by hunters (total number of records n = 77, of which 29 are from the North Caucasus, 24 from Azerbaijan, 13 from Armenia and 11 from

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Fig. 1. The Leopard, Panthera pardus, pictures taken by camera photo-traps in the Meghri Ridge of southern Armenia: adult male, 9 March 2005 (top) and sub-adult male, 8 February 2007 (bottom). Authors: I. KHOROZYAN, A. MALKHASYAN, M. BOYAJYAN.

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Georgia). The phrases “shot at the leopard”, “the leopard was wounded” and “the leopard was trapped” used by writers were considered as equivalent to killing.We also took into account the pictures taken by camera photo-traps (4 records and 10 pictures) and the results of faecal bile acid thin-layer chromatography as robust evidence for the presence of the Leopard (8 records, Armenia only) (B. LORTKIPANIDZE, pers. comm., BUTKHUZI 2004, KHOROZYAN & MALKHASYAN 2005, KHOROZYAN et al., in press). The compilation and analysis of Leopard killing records can sometimes be complicated by different presentations of the same event by different authors. In such cases, the authors bring their own subjective attitude which cannot now be separated from reality. Here is just one example of such information. ALEKPEROV (1947) wrote “…in November 1937, the Leopard tracks were observed from the mountain passes in front of Zakatali town”. BURCHAK-ABRAMOVICH & JAFAROV (1949) described this case as “in November 1937 the Leopard tracks were seen on the pass in front of Zakatali town” and added “…in December 1931 the Leopard tracks were seen in the headwaters of the Belokanchai River (Zakatali Nature Reserve) in the area of abandoned copper mines”. However, after many years, HEPTNER & SLUDSKY (1972) wrote “…in December 1931 it [the Leopard] was shot in the Zakatali Reserve and in November 1937 near the pass not far from Zakatali town”. In cases of such contradictions, we have relied on the earlier reported records.

History of the Leopard emergence in the Caucasus As molecular genetic studies show, the Leopard evolved as a distinct species ca 470,000825,000 years ago on the African continent (UPHYRKINA et al. 2001). Its Asian line was shaped later, from the late Pliocene to the early Pleistocene (ca 170,000-300,000 years ago), through migration from Africa to Asia via the Middle East. Then, between the middle and middle-late Pleistocene, the Leopard migrated from mainland Asia to the island of Java (MEIJAARD 2004). The northern boundary of its range in the Middle East lay in the Caucasus and the Kopetdag Mts in Turkmenistan, as no Leopard fossil remains have been found in the Pleistocene deposits of Eastern Europe (VERESHCHAGIN 1959). In the Caucasus, finds of Pleistocene Leopard bones are known from the Palaeolithic strata of the Kudaro I and III caves in the headwaters of the Rioni River in South Osetia, Georgia (VERESHCHAGIN 1959; two mandibles deposited in the Zoological Institute. St Petersburg, Russia), as well as from the Acheulian deposits (400,000-300,000 years ago) in the Azykh Cave of Karabakh (GADJIEV 2000). Holocene-dated fossils (10,000 B.C.) were discovered in the Sosruko Grotto, Baksan Canyon, Kabardino-Balkaria, Russia, and in the Dlinnokrylov Cave on the Araks River in Nakhichevan, Azerbaijan (VERESHCHAGIN 1959). Rock carvings depicting scenes of Leopards being hunted or captured by men dating from the late Stone Age (5000-4000 B.C.) and the early to middle Bronze Age (3000-2000 B.C.) were found on the Geghama Ridge and in Syunik Province of Armenia (MEZHLUMYAN 1985, KASABYAN & MANASERYAN 1998). The skull of a medium-sized Leopard dating from the early Bronze Age (3000 B.C.) was discovered in a grotto in the Akstafa district of north-western Azerbaijan (GADJIEV 2000). The absence of Leopard remains in the middle Pleistocene fauna of the Apsheron Peninsula, Azerbaijan, shows that the Leopard appeared first in the central Caucasus by migrating with the Armenian Mouflon Ovis orientalis gmelini and Indian Porcupine Hystrix indica from the south (VERESHCHAGIN 1959).

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Fig. 2. The sites of the most reliable Leopard records (killings, photo-trap pictures and scats biochemically proved to be of Leopard origin) in the Caucasus from the 1860s to 2007 (above) and current Leopard range in the Caucasus (below).

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In early historical times the Leopard was widespread in the Caucasus. Its range reached the plains of the North Caucasus, as is testified by the finds of bronze figurines of this carnivore in the Scythian tumuli of Ukraine and the Russian Plain (VERESHCHAGIN 1959). Up to the mid-19th century, the Leopard was common in the Caucasus and occurred in all habitats with vast forest tracts, dense scrublands, low-snow rocky massifs that were difficult of access, and, most important, with abundant resources of natural prey: Chamois, Rupicapra rupicapra, Bezoar Goat, Capra aegagrus, West Caucasian Tur, Capra caucasica, East Caucasian Tur, Capra cylindicornis, Roe Deer, Capreolus capreolus, and Wild Boar, Sus scrofa (DINNIK 1914, HEPTNER & SLUDSKY 1972, SLUDSKY 1973).

Habitats As our database shows, during the last two centuries the Leopard has preferred the most outof-the-way and the most remote mountainous habitats, in river headwaters covered by broadleaf and sparse forests, dry and mountain grasslands, subalpine and alpine meadows with numerous secluded rocky places and sufficient prey. An important factor is the availability of the low-snow territories which serve as overwintering sites for many ungulates and offer opportunities for successful hunting by the Leopard (HEPTNER & SLUDSKY 1972, KUDAKTIN 2002, ARABULI 2006). However, the Leopard also used to appear on the plains, e.g. in 1862 on the Colchic Lowland in Georgia, in 1896 and 1929-1932 on the Mugan Plain, in 1946 on the Apsheron Peninsula in Azerbaijan, and in 1954 in the Ararat Valley in Armenia (RADDE 1899, JAFAROV 1946, ALEKPEROV 1947, BURCHAK-ABRAMOVICH & JAFAROV 1949, VERESHCHAGIN 1959, HEPTNER & SLUDSKY 1972, SLUDSKY 1973). It possibly reached Kizliar town in the debouchment of the Terek River in Daghestan, Russia (HEPTNER & SLUDSKY 1972). There is information about Leopard visitations to the outskirts of big urban areas, e.g. in 1875 to Sochi town, in 1893 and 1940 to Tuapse town, in 1950 and 1956 to Lazarevskoe village (all in Krasnodar Territory, Russia); in 1923 and 1924 to Makhachkala town (Daghestan, Russia), in 1952 to Tskhinvali town in South Osetia, and in 1959-1960 to Akhaltsikhe town (all in Georgia), in 1937 to Zakatali town, in 1866, 1930, 1958-1959 to Lenkoran town (all in Azerbaijan) (RADDE 1899, HEPTNER & FORMOZOV 1941, ALEKPEROV 1947, VERESHCHAGIN 1959, KOTOV & RIABOV 1963, SLUDSKY 1973). Early records of this kind could reflect the much larger areas of forests and dense scrublands which in bygone times reached as far as the settlements (HEPTNER & SLUDSKY 1972). Later records could result from the shortage of natural prey that forced the Leopards to hunt near humans on livestock and dogs (KOTOV & RIABOV 1963).

Scales of eradication In total, during the period from 1861 to 2007, the Leopard death toll in the Caucasus was 118 individuals, including 44 in Azerbaijan, 38 in the North Caucasus of Russia, 18 in Armenia and 18 in Georgia. Up to the early 1970s, the Leopard and other large carnivores were ruthlessly wiped out as vermin. In a number of areas, the bounties paid by the government accelerated extermination and led to the extinction of the Leopard in most of its former range (DINNIK 1914). The dynamics of Leopard eradication in the Caucasus during this period are shown in Fig. 4. The peaks of the killing curves were reached in the periods 1880-1910 (17 Leopards shot) and 1920-1960 (18) in the North Caucasus, in 1950-1960 (7) in Georgia, and

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in 1940-1970 in Azerbaijan and Armenia (22 and 12, respectively). In the text below, we will consider the history of Leopard survival in each of four countries of the Caucasus in more detail. We recognise that the statistical data on Leopard killings tend to underestimate the real scale of its extermination. The reason is that not many cases remain unknown, especially since the early 1970s when the Leopard was declared an officially protected mammal in all Caucasus countries (then the Soviet Union republics) and poaching episodes have been hushed up in order to avoid punishment.

North Caucasus of Russia The “Kubanskaya okhota” (Kuban Hunt) wildlife sanctuary of the Grand Duka Sergey Mikhailovich was founded in 1882 in the western part of the North Caucasus (south of today’s Krasnodar Territory and a southern portion of Adygea). In 1924 it was reorganised into the Kavkazsky (Caucasus) Nature Reserve (AKATOV et al. 1990). The Leopard was perceived as vermin, destroying the traditionally hunted species, and it was relentlessly pursued by the large staff of rangers who could earn 25 rubles per skin (DINNIK 1914). As a result, during the period of existence of the “Kubanskaya okhota” (until 1909), sixteen Leopards were killed in the Kisha, Umpyr, Laba, Shisha and Malaya Laba river canyons, in Nikitskaya Balka and Pshekish localities. Poisoned baits (meat with strychnine) and steel traps mostly left near the stream fords were widely used. Of the 16 Leopards killed, seven were shot, seven poisoned and two trapped (NASIMOVICH 1941, HEPTNER & SLUDSKY 1972, SLUDSKY 1973). The depressing outcome was the almost complete disappearance of the Leopard from this area, and even the establishment of the Caucasus Nature Reserve failed to change the situation. Rumours of sightings of the Leopard itself or its tracks or hearing its grumbling sounds continued to come in, but not a single animal was harvested. However, in 1936 in the Riazan district of Krasnodar Territory, a Leopard killed a local man and attacked several people and animals until it was cornered and wounded (KOTOV & RIABOV 1963). GINEEV (1983) and KUDAKTIN (1985) indicate that a Leopard was wounded in winter 1972 in the Artagarych locality on the headwaters of the Pshekha River in Krasnodar Territory. It is most likely that this individual was a vagrant, as the northern slopes of the Great Caucasus Ridge have been used by Leopards, mainly in the lowest eastern part of this ridge in Daghestan (KUDAKTIN 1985). Meanwhile, up to the mid-1950s, the Leopard continued to survive on the Black Sea slope of the Great Caucasus Ridge, in Karachaevo-Cherkessia and the Arkhyz district of the Teberdinsky Nature Reserve. The facts are the following: in 1929, two Leopards were shot in Orekhovaya Poliana near the Sochi River; in summer 1940, one individual was killed close to the “Pervoe maya” sanatorium, not far from Tuapse town; two Leopards were harvested in the Urup River gorge – a male in 1937 and a female in December 1940; in 1948, a male was trapped between the Laba and Urup rivers in Karachaevo-Cherkessia; in 1956, a Leopard frequented Lazarevskoe village to prey on dogs until it was shot (NASIMOVICH 1941, KOTOV & RIABOV 1963, PENZIKOV 1986). Since the mid-1950s, no direct evidence (such as skins) for the existence of the Leopard in the western North Caucasus is known, but there are some records of indirect presence signs (tracks and roar) and even observations (KUDAKTIN 1985). In 1996 and 2001, large tracks which might belong to the Leopard were found in the Caucasus Nature Reserve (LUKAREVSKY et al. 2004).

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From the 1920s, data on Leopard harvests began to come from the central and eastern parts of the North Caucasus: in the 1920s two Leopards were shot in North Osetia; in spring 1924, one predator was destroyed near Agach-aul village close to Makhachkala town, Daghestan; in the 1940-1950s the Leopard was killed near Lars, North Osetia; in 1949 and 1953, two individuals were harvested in the Sunzha River canyon, Checheno-Ingushetia; in 1957 and 1958, two Leopards were shot in the Chegem River canyon, Kabardino-Balkaria (HEPTNER & FORMOZOV 1941, VERESHCHAGIN 1959, HEPTNER & SLUDSKY 1972, GINEEV 1983, AIUNTS & SHALYBKOV 1990). From 1980 onwards, only two cases of Leopard killings are known throughout the entire North Caucasus: in 1981 an adult male was shot in the Gumbetovsky district of Daghestan, and in April 2001 a female accompanied by two 2-month-old male cubs was killed close to the border between Kabardino-Balkaria and Georgia (the cubs were transferred to the Novosibirsk Zoo where they are still living) (SPASSKAYA & SAIDALIEVA 1983, LUKAREVSKY et al. 2004, RAFFEL 2004). LUKAREVSKY (2006) thought that the Leopard could survive in the basin of the Avarskoe Koisu River in Daghestan and, possibly, in the Erzi Nature Reserve of Ingushetia.

Georgia The greatest numbers of Leopard trophies in Georgia were taken in two places: in 18891999, three Leopards were poisoned in Borjomi, and in winter 1959-1960 three individuals were killed near Akhaltsikhe town (RADDE 1899, HEPTNER & SLUDSKY 1972). Single individuals were also killed in the canyon of the Bzyb River in Abkhazia in 1861, in the canyon of the Sken River in the Colchic Lowlands in 1862, in Lagodekhi Reserve in 1914, in South Osetia in 1950 and 1952, in the Zedazen Forest of Saguramo Nature Reserve in 1954, and in the Assa River basin in 1979 and 1986 (RADDE 1899, VERESHCHAGIN 1959, HEPTNER & SLUDSKY 1972, SLUDSKY 1973, CHIKOVANI et al. 1990, ARABULI 2006). The last known Leopard trophy is an adult male that was shot in the Arkhoti River basin in north-eastern Georgia in 1999 (ARABULI 2006). In 2004-2005, seven pictures of a young male were taken by camera photo-traps in the Vashlovani Nature Reserve in the extreme south-eastern part of the country (BUTKHUZI 2004, B. LORTKIPANIDZE, pers. comm.). LUKAREVSKY (2006) suggests that the Leopard can penetrate as far as Tusheti, a highland area on the border with Russia’s North Caucasus, but suitable habitats are very limited and are unlikely to ensure population survival. ARABULI (2006) claims that the Leopard also exists in south-western Georgia, Adjaria, but the quality of the only known record is not convincing (observation of an animal from a distance of 3 km).

Azerbaijan The Talysh Mts located in the extreme south-eastern corner occupy the top position for Leopard trophies harvested in this country. Out of 44 Leopards killed in Azerbaijan, 30 came from these mountains. Moreover, several individuals were shot in Karabakh, Nakhichevan and on the Great Caucasus Ridge. The maximum intensity of Leopard extermination was recorded in the period from 1940 to 1970 during which 22 animals were killed: in 1941, 1959 and 1967, three predators were shot in the Shusha, Mardakert and Gadrut districts of Karabakh; in 1944, 1946, winter of 1958-1959, 1960, 1961 and 1969, thirteen individuals were killed in the Talysh Mts near Lenkoran and Astara towns; in 1946, two Leopards were

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killed in Nakhichevan to the east of Ordubad town, one male was shot in Bilga village on the Apsheron Peninsula, and one individual was harvested near Yantak village in the Kelbajar district between Armenia and Karabakh; in 1952, one Leopard was killed near Negram railway station in the Araks River canyon and another individual was shot in 1958 in the Geidash (or Gaidag) Canyon, both in Nakhichevan (JAFAROV 1946, ALEKPEROV 1947, BURCHAK-ABRAMOVICH & JAFAROV 1949, ALEKPEROV 1966, HEPTNER & SLUDSKY 1972, SLUDSKY 1973). In 1975, the Leopard was harvested in the Kakhsky district in the north-west of the country, the last record on the Azerbaijani part of the Great Caucasus Ridge (GADJIEV & NASIBOV 1979). Since then, three Leopards are known to have been shot: one in 2000 near the Kility border post in Nakhichevan and two in 2002 in the Talysh Mts (ASKEROV 2002, pers. comm.). The last known Leopard trophy in Azerbaijan is dated 27 November 2006; it was a male shot by locals near Akhtakhana village in the Jalilabad district to the north of the Talysh Mts (SULEIMANOVA 2006). On 24 February 2007, a leopard was captured by phototrap in the Talysh Mts (E. ASKEROV, pers. comm.). Up to the mid-1950s, anecdotal Leopard records also used to be received from the Zakatali Nature Reserve and its vicinity in the extreme north-west, but since then there has been no information from this area.

Armenia Unlike other regions of the Caucasus, Leopard records in Armenia only began to come in the late 1930s. Here, most of the trophies were harvested in the area of the Khosrov Nature Reserve (10 of 17 specimens). The maximum number (12 Leopards) was harvested in 19401970: before 1946, three individuals with no precise localities indicated; in 1956, 1959, 1960 and 1968, six predators were shot in Khosrov Nature Reserve; in 1953 and 1959, two Leopards were killed near the villages of Bartsruni and Zaritap in the Azizbekov (now Vayk) district; in 1954, one Leopard was shot near Dvin village in the Artashat district of the Ararat Valley (skins No. 6435 and 6437 and stuffed specimens 6436 and 6438 in the Institute of Zoology, Yerevan; most likely these are the individuals mentioned by BURCHAKABRAMOVICH & JAFAROV 1949; stuffed specimens No. 241.1 and 241.2 in the Museum of Nature, Yerevan; HEPTNER & SLUDSKY 1972, SLUDSKY 1973, GASPARYAN & AGADJANYAN 1974, M. ADAMYAN and V. HAKOPYAN, pers. comm.). Before the early 1970s, the Leopard also occurred in northern Armenia (DAL 1954, GASPARYAN & AGADJANYAN 1974, KASABYAN 2001), but since then it has become extinct there (KHOROZYAN & ABRAMOV 2005, KHOROZYAN et al. 2005). Information from our database differs from that published earlier by GASPARYAN & AGADJANYAN (1974) and AIRUMYAN & GASPARYAN (1976). According to these authors, 73 Leopards were harvested in Armenia in the period 1952-1971, of which 54 came from just the 4 years 1967-1971, with a maximum of 16 skins in 1971. Almost half of all Leopards were killed in northern Armenia and so the authors postulated a northward expansion of the Leopard population in Armenia. Before that, 9 Leopards were shot in 1952-1958 and 8 in 19591966 (AIRUMYAN & GASPARYAN 1976; but 12 in 1958-1966, GASPARYAN & AGADJANYAN 1974). Setting a rather high bounty in 1959 did not affect the dynamics of eradication. These publications were based on information from the national skin supply centres and it is most likely that the majority of the skins belonged to the Lynx which is common in Armenia. In 1972 the Leopard was declared an officially protected mammal in Armenia and was listed in the national Red Data Book as “endangered” (GASPARYAN & AGADJANYAN 1974).

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%

North Caucasus

60

Georgia Azerbaijan Armenia

40

20

0 killings

observations

tracks, roar, prey remains, scats, scrapes, dens

attacks on humans, livestock and pets

Fig. 3. Distribution of the types of Leopard records in the Caucasus from the 1860s to the present.

The last records of Leopard killings in Armenia were obtained from the Bargushat Ridge (adult male, November 1997), Garni district of Khosrov Nature Reserve (in 2000), and the disputed lands adjoining the extreme south-east of Armenia and controlled by Armenian troops (Meghri Ridge, adult male, winter 2005-2006). In addition, in 2004-2007, the presence of the Leopard in the eastern Khosrov Reserve and on the Meghri Ridge was unambiguously proved by camera photo-trapping and by the determination of the Leopard origin of scats by faecal bile acid thin-layer chromatography (KHOROZYAN & MALKHASYAN 2005, KHOROZYAN et al. in press; Fig. 1). In 2005-2006, Leopard scats were also found on the Zangezour Ridge (unpubl. data).

Conclusion All recent and current Leopard records in the Caucasus are confined to the following regions: central (Chegem River canyon, Kabardino-Balkaria) and eastern (headwaters of the Andiiskoe Koisu and Avarskoe Koisu rivers in Daghestan and the Erzi Nature Reserve in Ingushetia) parts of the North Caucasus; Vashlovani Nature Reserve and north-eastern part of Georgia; Talysh Mts and Zangezour Ridge in Nakhichevan, Azerbaijan; Khosrov Nature Reserve, Zangezour, Bargushat and Meghri Ridges in Armenia (Fig. 2). The status of the Leopard in Karabakh is uncertain because of the unsettled military and political conflict between Armenia and Azerbaijan over this territory which prevents the acquisition of reliable information and the implementation of research. Also, because of the political instability in the Chechn Republic, the Leopard cannot be studied in this republic and in the adjacent areas of Ingushetia and Daghestan (LUKAREVSKY 2006). The areas of Kabardino-Balkaria, Ingushetia, Daghestan and the Vashlovani Nature Reserve in Georgia are supposedly inter-connected, and some authors believe in the possible existence of an independent population in this region (GINEEV 1983, LUKAREVSKY et al.

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14

North Caucasus Georgia Azerbaijan Armenia

numbers of trophies

12 10 8 6 4 2

2000-2007

1990-2000

1980-1990

1970-1980

1960-1970

1950-1960

1940-1950

1930-1940

1920-1930

1910-1920

1900-1910

1890-1900

1880-1890

1870-1880

1860-1870

0

Fig. 4. The Leopard numbers harvested in the Caucasus from the 1860s to the present.

2004). However, a more plausible option would be the connection of this region with southern Armenia through the Karabakh Highland and the Mrovdag Ridge (to the south-east of Lake Sevan) (Fig. 2). The implementation of in-depth and systematic research in southeastern Georgia and north-western Azerbaijan could shed light on this issue. Meanwhile, some scientists believe that the Daghestan portion of the range is more closely linked with Azerbaijan than with the central part of the North Caucasus (ALEKEROV 1947, HEPTNER & FORMOZOV 1941). The Talysh Mts are isolated from the rest of the Leopard range in the Caucasus, but are directly linked with Iran. In their turn, the Khosrov Nature Reserve, Zangezour, Bargushat and Meghri ridges in Armenia and the Nakhichevan part of the Leopard range are connected by the common Zangezour Ridge extending along the Armenian-Azerbaijani (Nakhichevan Republic) state border. The Leopard populations are resident in southern Armenia and the Talysh Mts as they are based on territorial adult individuals who mark their home ranges by scrapes and scats (LUKAREVSKY et al. 2004, KHOROZYAN & ABRAMOV 2005, KHOROZYAN & MALKHASYAN 2005). However, they are too small for independent long-term survival and must depend, at least partially, on the immigration of new founders from the much larger population in adjacent north-western Iran (KIABI et al. 2002).

Acknowledgements. This publication is part of the first author’s PhD dissertation at the Zoological Institute in St Petersburg. We would like to thank N. U. MANASERYAN and M. S. ADAMYAN (Institute of Zoology, Armenia) for assistance during the work with their museum collections and B. LORTKIPANIDZE (NACRES, Georgia), A. SAVELJEV (Institute of Game Management and Fur Farming, Russia), M. RAFFEL (Allwetterzoo Münster, Germany), E. ASKEROV (WWF, Azerbaijan) and V. HAKOPYAN (Institute of Zoology, Armenia) for

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information support. Financial support of the Leopard research projects in Armenia, the results of which are used in this study, has been generously provided by the People’s Trust for Endangered Species (UK) and the Panthera/WCS Kaplan Award Program (USA).

References AIRUMYAN, K. A. & K. M. GASPARYAN (1976): Rare ungulates and carnivores of Armenia [in Russian]. p. 35–43. In: V. E. SOKOLOV (Ed.), Redkie Mlekopitayuschie Fauny SSSR. – Nauka, Moscow. AIUNTS, K. P. & A. M. SHALYBKOV (1990): Kabardino-Balkarian Nature Reserve [in Russian]. p. 35– 45. In: V. E. SOKOLOV & E. E. SYROECHKOVSKY (Eds.), Zapovedniki Kavkaza. – Mysl, Moscow. AKATOV, V. V., K. YU. GOLGOFSKAYA, L. G. GORCHARUK, V. V. DUROV, V. V. KIPIANI, A. N. KUDAKTIN, A. S. NEMTSEV, M. V. PRIDNIA, P. A. TILBA & V. G. TOPILINA (1990): Caucasus Nature Reserve [in Russian]. p. 69–101. In: V. E. SOKOLOV & E. E. SYROECHKOVSKY (Eds.), Zapovedniki Kavkaza. – Mysl, Moscow. ALEKPEROV, A. M. (1947): New data about the South Caucasus Leopard (Panthera pardus tullianus Valenc.) [in Russian]. – Doklady Akademii Nauk Azerbaidzhanskoi SSR 3: 126–128. ALEKPEROV, Kh. M. (1966): Mammals of South-Western Azerbaijan [in Russian]. – Izdatelstvo Akademii Nauk AzSSR, Baku. ARABULI, A. B. (2006): On distribution of the leopard (Panthera pardus ciscaucasica) in South Caucasus within Georgia [in Russian]. – Zoologicheskii zhurnal 85: 910–912. ASKEROV, E. K. (2002): New data on leopard (Panthera pardus L.) in Azerbaijan and measures for its conservation [in Russian]. – Bilgi (Baku) 3: 69–72. BURCHAK-ABRAMOVICH, N. O. & R. D. JAFAROV (1949): The leopard (Panthera pardus tullianus Valenc.) on Apsheron Peninsula [in Russian]. – Trudy Estestvenno-istoricheskogo Muzeya Akademii Nauk Azerbaidzhanskoi SSR 3: 86–104. BUTKHUZI, L. (2004): Breaking news - leopard in Georgia. – Caucasus Environment 2: 49–51. CHIKOVANI, T. G., N. V. VRONSKY, B. E. KURASHVILI & Z. D. SIRAKHULIDZE (1990): SaguramoMariamjvarian Nature Reserve [in Russian]. p. 191–200. In: V. E. SOKOLOV & E. E. SYROECHKOVSKY (Eds.), Zapovedniki Kavkaza. – Mysl, Moscow. DAL, S. K. (1954): Animal World of Armenian SSR. Vol. 1. Vertebrates [in Russian]. – Izdatelstvo Akademii Nauk Arm SSR, Yerevan. DINNIK, N. Ya. (1914): Mammals of the Caucasus. Part II. Carnivores [in Russian]. - Tipografiya K.P. Kozlovskogo, Tiflis. GADJIEV, D. V. (2000): Leopard - Panthera pardus L. [in Russian]. p. 577–578. In: T. KERIMLY (Ed.), Zhivotnyi Mir Azerbaijana. Vol. 3 [in Russian]. - Elm, Baku. GADJIEV, D. V. & S. B. NASIBOV (1979): Wild cats of Azerbaijan. [in Russian]. p. 251–253. In: V. E. SOKOLOV (Ed.), Ekologicheskie Osnovy Okhrany i Ratsionalnogo Ispolzovaniya Khischnykh Mlekopitayuschikh. - Nauka, Moscow. GASPARYAN, K. M. & F. S. AGADJANYAN (1974): On leopard in Armenia [in Russian]. Biologicheskii zhurnal Armenii 27: 84–88. GINEEV, A. M. (1983): Conservation issues of the Caucasian leopard [in Russian]. p. 96–97. In: V. E. SOKOLOV (Ed.), Redkie Vidy Mlekopitayuschikh SSSR i Ikh Okhrana. - Izdatelstvo Akademii Nauk SSSR, Moscow. GRID (2002): Caucasus Environmental Outlook, CEO-2002. – GRID, Tbilisi. HEPTNER, V. G. & A. N. FORMOZOV (1941): Mammals of Daghestan [in Russian]. p. 3–74. In: N. N. FILIPPOV (Ed.), Sbornik Trudov Gosudarstvennogo Zoologicheskogo Muzeya MGU. – KOIZ, Moscow. HEPTNER, V. G. & A. A. SLUDSKY (1972): Mammals of the Soviet Union. Vol.2. Part 2. Carnivores (Hyenas and Cats) [in Russian]. - Vysshaya Shkola, Moscow.

Mammalia

23

JAFAROV, R. D. (1946): The leopard (Felis pardus L.) on Apsheron [in Russian]. - Izvestiya Akademii Nauk Azerbaidzhanskoi SSR, otdelenie geologo-khimicheskikh nauk i nefti (Baku) 2: 106-107. KASABYAN, M. G. (2001): Carnivorous Mammals of Armenia [in Russian]. – PhD dissertation, Institute of Zoology, Yerevan. KASABYAN, M. G. & N. U. MANASERYAN (1998): Carnivorous mammals in the Holocene of Armenia [in Russian]. - Materialy respublikanskoi nauchnoi konferentsii po zoologii 14-15 maya 1998 (Yerevan), p. 63–64. KHOROZYAN, I. G. & A. V. ABRAMOV (2005): The spatial structure and conservation of the Persian leopard population in Armenia [in Russian]. p. 199–202. In: V. V. ROZHNOV & F. A. TEMBOTOVA (Eds.), Mlekopitayuschie Gornykh Territorii. – Tovarischestvo nauchnykh izdanii KMK, Moscow. KHOROZYAN, I. G., G. F. BARYSHNIKOV & A. V. ABRAMOV (2006): Taxonomic status of the leopard, Panthera pardus (Carnivora, Felidae) in the Caucasus and adjacent areas. – Russian Journal of Theriology 5: 41–52. KHOROZYAN, I. G., A. CAZON, A. G. MALKHASYAN & A. V. ABRAMOV (in press): Using fecal bile acid thin-layer chromatography to study the leopard Panthera pardus population. – Biology Bulletin, Moscow. KHOROZYAN, I. & A. MALKHASYAN (2005): Persian leopard photographed in Armenia. - Cat News (Bougy) 43: 13. KHOROZYAN, I., A. MALKHASYAN & S. ASMARYAN (2005): The Persian leopard prowls its way to survival. - Endangered Species Update (Ann Arbor) 22: 51–60. KIABI, B. H., B. F. DARESHOURI, R. A. GHAEMI & M. JAHANSHAHI (2002): Population status of the Persian leopard (Panthera pardus saxicolor Pocock, 1927) in Iran. – Zoology in the Middle East 26: 41–47. KOTOV, V. A. & L. S. RIABOV (1963): The commercial and valuable mammals of the foothill and mountainous areas of Krasnodar Territory, Moscow. – Trudy Kavkazskogo gosudarstvennogo zapovednika (Moscow) 7: 3–226. KUDAKTIN, A. N. (1985): Leopard in the Caucasus [in Russian]. – Okhota i okhotnichie khoziaistvo (Moscow) 11: 16. KUDAKTIN, A. N. (2002): Lynx and leopard in protected areas of the Caucasus [in Russian]. p. 96–103. In: A. S. ZHELTUKHIN & V. V. KOCHETKOV (Eds.), Byulleten Nauchno-Metodicheskogo Tsentra po Izucheniyu Krupnykh Khischnykh Mlekopitayuschikh v Zapovednikakh Rossii. Vyp. 2. – Ministerstvo prirodnykh resursov Rossii, Moscow. LUKAREVSKY, V. S. (2006): The spotted hostage. Home for the leopard [in Russian]. – Panda Times May (Moscow) 2006: 8–9. LUKAREVSKY, V., E. ASKEROV & G. HAZARYAN (2004): Condition of the leopard population in the Caucasus. – Beiträge zur Jagd- und Wildforschung 29: 305–319. MEIJAARD, E. (2004): Biogeographic history of the Javan leopard Panthera pardus based on a craniometric analysis. – Journal of Mammalogy 85: 302–310. MEZHLUMYAN, S. K. (1985): Hunting on the rock carvings in the Geghama Mountains and Syunik [in Russian]. – Biologicheskii zhurnal Armenii (Yerevan) 38: 685–696. NASIMOVICH, A. (1941): Leopard in western Caucasus [in Russian]. – Priroda i sotsialisticheskoe khoziaistvo (Moscow) 8: 260–265. PENZIKOV, Yu. A. (1986): The last leopards in Karachaevo-Cherkessia [in Russian]. – Priroda (Moscow) 3: 96. RADDE, G. I. (1899): Collection of the Caucasus Museum. Vol. 1. Zoology [in Russian]. - Tipografiya Kantseliarii Glavnokomanduyuschego grazhdanskoi chastiu na Kavkaze (Tiflis). RAFFEL, M. (2004): Persian Leopard (Panthera pardus saxicolor). – EEP Annual Report 2004 Allwetter Zoo Münster. SLUDSKY, A. A. (1973): Distribution and abundance of wild cats in the USSR [in Russian]. – Trudy Instituta zoologii Akademii Nauk Kazakhskoi SSR (Alma-Ata) 34: 5–106. SPASSKAYA, T. & M. Saidalieva (1983): Leopard in Daghestan [in Russian]. – Okhota i okhotnichie khoziaistvo (Moscow) 12: 10.

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SULEIMANOVA, G. (2006): A “king” of Azerbaijani forests recently killed [in Russian]. – Internetgazeta “Zerkalo” 2 December 2006 UPHYRKINA, O., W. E. JOHNSON, H. QUIGLEY, D. MIQUELLE, L. MARKER, M. BUSH & S. J. O’BRIEN (2001): Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus. – Molecular Ecology (Loughborough) 10: 2617–2633. VERESHCHAGIN, N. K. (1959): Mammals of the Caucasus. History of Fauna Formation [in Russian]. – Izdatelstvo Akademii Nauk SSSR (Moscow & Leningrad).

Authors’ address: Igor G. Khorozyan and Alexei V. Abramov, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, St Petersburg 199034, Russia. – Contact c/o I. G. Khorozyan, Mashtotz Ave. 48/17, Yerevan 375009, Armenia. E-mail: [email protected].