Caspian tiger

The Caspian tiger was a tiger from a specific population of the Panthera tigris tigris subspecies that was native to the eastern Turkey, northern Iran, Mesopotamia, the Caucasus around the Caspian Sea through Central Asia to northern Afghanistan and Xinjiang in western China.[5] It inhabited sparse forests and riverine corridors in this region until the 1970s.[1] This population was assessed as extinct in 2003.[2]

Caspian tiger
Tiger from the Caucasus in Berlin Zoo, 1899[1]

Extinct  (~ ca.1970-2003)  (IUCN 3.1)[2]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species:
P. tigris
Subspecies:
†P. t. tigris
Trinomial name
Panthera tigris tigris
Original distribution of the Caspian tiger (in black)
Synonyms[3][4]
  • P. t. virgata (Illiger, 1815)
  • P. t. lecoqi
  • P. t. septentrionalis
  • P. t. trabata

Felis virgata was the scientific name proposed in 1815 by Johann Karl Wilhelm Illiger for tigers in the Caspian Sea area.[6] It was traditionally recognised as a distinct subspecies, Panthera tigris virgata. However, results of phylogeographic analysis indicate that the Caspian and Siberian tiger populations shared a common continuous geographic distribution until the early 19th century.[7]

Some Caspian tiger individuals were intermediate in size between Siberian and Bengal tigers.[8][3][9]

The Caspian tiger was also called Balkhash tiger, Hyrcanian tiger, Turanian tiger,[2] and Babre Mazandaran (Persian: ببرِ مازندران‎), depending on the region of its occurrence.[10]

Taxonomy

Felis virgata was the scientific name used by Illiger in 1815 when he described the greyish tigers in the area of the Caspian Sea and Persia.[6] In 1929, Reginald Innes Pocock subordinated the tiger to the genus Panthera.[11] For several decades, the Caspian tiger was considered a distinct tiger subspecies.[3][4]

In 1999, the validity of several tiger subspecies was questioned. Most putative subspecies described in the 19th and 20th centuries were distinguished on basis of fur length and colouration, striping patterns and body size, hence characteristics that vary widely within populations. Morphologically, tigers from different regions vary little, and gene flow between populations in those regions is considered to have been possible during the Pleistocene. Therefore, it was proposed to recognize only two tiger subspecies as valid, namely P. t. tigris in mainland Asia, and P. t. sondaica in the Greater Sunda Islands and possibly in Sundaland.[12]

At the start of the 21st century, genetic studies were carried out using 20 tiger bone and tissue samples from museum collections and sequencing at least one segment of five mitochondrial genes. Results revealed a low amount of variability in the mitochondrial DNA in Caspian tigers; and that Caspian and Siberian tigers were remarkably similar, indicating that the Siberian tiger is the genetically closest living relative of the Caspian tiger. Phylogeographic analysis indicates that the common ancestor of Caspian and Siberian tigers colonized Central Asia via the Gansu−Silk Road region from eastern China less than 10,000 years ago, and subsequently traversed eastward to establish the Siberian tiger population in the Russian Far East. The Caspian and Siberian tigers were likely a single contiguous population until the early 19th century, but became isolated from another due to fragmentation and loss of habitat during the Industrial Revolution.[7]

In 2015, morphological, ecological and molecular traits of all putative tiger subspecies were analysed in a combined approach. Results support distinction of the two evolutionary groups continental and Sunda tigers. The authors proposed recognition of only two subspecies, namely P. t. tigris comprising the Bengal, Malayan, Indochinese, South Chinese, Siberian and Caspian tiger populations, and P. t. sondaica comprising the Javan, Bali and Sumatran tiger populations. Tigers in mainland Asia fall into two clades, namely a northern clade formed by the Caspian and Siberian tiger populations, and a southern clade formed by populations in remaining mainland Asia.[13]

In 2017, the Cat Specialist Group revised felid taxonomy and now recognizes the tiger populations in continental Asia as P. t. tigris.[5] However, a genetic study published in 2018 supported six monophyletic clades, with the Amur and Caspian tigers being distinct from other mainland Asian populations, thus supporting the traditional concept of six living subspecies.[14]

Characteristics
Skin of a Caspian tiger from Iran
Comparative illustration of the stripe patterns on the tails of Caspian (left) and Siberian tigers (right)[8]
Illustration of two Caspian tigers

Fur

Photographs of skins of Caspian and Amur tigers indicate that the main background colour of the Caspian tiger's fur varied and was generally brighter and more uniform than that of the Siberian tiger. The stripes were narrower, fuller and more closely set than those of tigers from Manchuria. The colour of its stripes was a mixture of brown or cinnamon shades. Pure black patterns were invariably found only on head, neck, the middle of the back and at the tip of the tail. Angular patterns at the base of the tail were less developed than those of Far Eastern populations. The contrast between the summer and winter coats was sharp, though not to the same extent as in Far Eastern populations. The winter coat was paler, with less distinct patterns. The summer coat had a similar density and hair length to that of the Bengal tiger, though its stripes were usually narrower, longer and closer set. It had the thickest fur amongst tigers, possibly due its occurrence in the temperate parts of Eurasia.[8][3][9]

Size

The Caspian tiger ranked among the largest cats that ever existed.[3][8][15] Males had a body length of 270–295 cm (106–116 in) and weighed 170–240 kg (370–530 lb); females measured 240–260 cm (94–102 in) in head-to-body and weighed 85–135 kg (187–298 lb).[3] Maximum skull length in males was 297–365.8 mm (11.69–14.40 in), while that of females was 195.7–255.5 mm (7.70–10.06 in).[8] Its occiput was broader than of the Bengal tiger.[12]

Some individuals attained exceptional sizes. In 1954, a tiger was killed near the Sumbar River in Kopet-Dag whose stuffed skin was put on display in a museum in Ashgabat. Its head-to-body length was 2.25 m (7.4 ft). Its skull had a condylobasal length of about 305 mm (12.0 in), and zygomatic width of 205 mm (8.1 in). Its skull length was 385 mm (15.2 in), hence more than the known maximum of 365.8 mm (14.40 in) for this population, and slightly exceeding skull length of most Siberian tigers.[8] In Prishibinske, a tiger was killed in February 1899. Measurements after skinning revealed a body length of 270 cm (8.9 ft) between the pegs, plus a 90 cm (3.0 ft) long tail, giving it a total length of about 360 cm (11.8 ft). Measurements between the pegs of up to 2.95 m (9.7 ft) is known.[3] According to Satunin it was "a tiger of immense proportions" and "no smaller than the common Tuzemna horse." It had rather long fur.[8]

Skull size and shape of Caspian tigers significantly overlap with and are almost indistinguishable from other tiger specimens in mainland Asia.[16]

Distribution and habitat
Shore of the Türkmenbaşy Gulf at the Caspian Sea
The Tigris River outside Mosul in Iraq
Landscape in the Altai Mountains

Historical records show that the distribution of the tiger in the region of the Caspian Sea was not continuous but patchy, and associated with wetlands such as river basins, lake edges and sea shores.[8] In the 19th century, tigers occurred in:

Its former distribution can be approximated by examining the distribution of ungulates in the region.[23] Wild boar was the numerically dominant ungulate occurring in forested habitats, along watercourses, in reed beds and in thickets of the Caspian and Aral Seas. Where watercourses penetrated deep into desert areas, suitable wild pig and tiger habitat was often linear, only a few kilometers wide at most. Red and roe deer occurred in forests around the Black Sea to the western side and around the southern side of the Caspian Sea in a narrow belt of forest cover. Roe deer occurred in forested areas south of Lake Balkash. Bactrian deer occurred in the narrow belt of forest habitat on the southern border of the Aral Sea, and southward along the Syr-Darya and Amu-Darya rivers.[8]

Local extinction
Tiger killed in northern Iran, early 1940s

The demise of the Caspian tiger began with the Russian colonisation of Turkestan during the late 19th century.[24] Its extirpation was caused by several factors:

  • Tigers were killed by large parties of sportsmen and military personnel who also hunted tiger prey species such as wild pigs. The range of wild pigs underwent a rapid decline between the middle of the 19th century and the 1930s due to overhunting, natural disasters, and diseases such as swine fever and foot-and-mouth disease, which caused large and rapid die-offs.[8]
  • The extensive reedbeds of tiger habitat were increasingly converted to cropland for planting cotton and other crops that grew well in the rich silt along rivers.[24]
  • Tigers were already vulnerable due to the restricted nature of their distribution, having been confined to watercourses within the large expanses of desert environment.[23]

Until the early 20th century, the regular Russian army was used to clear predators from forests, around settlements, and potential agricultural lands. Until World War I, about 100 tigers were killed in the forests of Amu-Darya and Piandj Rivers each year. High incentives were paid for tiger skins up to 1929. The prey base of tigers, wild pigs and deer, were decimated by deforestation and subsistence hunting by the increasing human population along the rivers, supported by growing agricultural developments.[25] By 1910, cotton plants were estimated to occupy nearly one-fifth of Turkestan's arable land, with about one half located in the Fergana Valley.[26]

Last sightings

In Iraq, a tiger was killed near Mosul in 1887.[1][17]

In Georgia, the last known tiger was killed in 1922 near Tbilisi, after taking domestic livestock.[1][27] Its stuffed body was put on display in the Georgian National Museum.[28][29]

In China, tigers disappeared from the Tarim River basin in Xinjiang in the 1920s.[1][27] From the Manasi River basin in the Tian Shan Range west of Ürümqi they reportedly disappeared in the 1960s.[8]

In Turkey, a pair of tigers was allegedly killed in the area of Selçuk in 1943.[30] Several tiger skins found in the early 1970s near Uludere indicated the presence of a tiger population in eastern Turkey.[31][32] Questionnaire surveys conducted in this region revealed that one to eight tigers were killed each year until the mid-1980s, and that tigers likely had survived in the region until the early 1990s. Due to lack of interest, in addition to security and safety reasons, no further field surveys were carried out in the area.[18]

In Kazakhstan, the last Caspian tiger was recorded in 1948, in the environs of the Ili River, the last known stronghold in the region of Lake Balkhash.[8]

In Iran, one of the last known tigers was shot in Golestan National Park in 1953. Another individual was sighted in the Golestan area in 1958.[9]

In Turkmenistan, the last known tiger was killed in January 1954 in the Sumbar River valley in the Kopet-Dag Range.[33] The last record from the lower reaches of the Amu-Darya river was an unconfirmed observation in 1968 near Nukus in the Aral Sea area. By the early 1970s, tigers disappeared from the river's lower reaches and the Pyzandh Valley in the Turkmen-Uzbek-Afghan border region.[8]

The Piandj River area between Afghanistan and Tajikistan was a stronghold of the Caspian tiger until the late 1960s. The latest sighting of a tiger in the Afghan-Tajik border area dates to 1998 in the Babatag Range.[25]

Behaviour and ecology
Mosaic of an elephant attacking a tiger, from Roman Syria, which occupied parts of what is now Anatolia and Mesopotamia[34]

No information is available for home ranges of Caspian tigers. In search for prey, they possibly prowled widely and followed migratory ungulates from one pasture to another. Wild pigs and cervids probably formed their main prey base. In many regions of Central Asia, Bactrian deer and roe deer were important prey species, as well as Caucasian red deer, goitered gazelle in Iran; Eurasian golden jackals, jungle cats, locusts, and other small mammals in the lower Amu-Darya River area; saiga, wild horses, Persian onagers in Miankaleh peninsula; Turkmenian kulans, Mongolian wild asses, and mountain sheep in the Zhana-Darya and around the Aral Sea; Manchurian wapiti and moose in the area of Lake Baikal. They caught fish in flooded areas and irrigation channels. In winter, they frequently attacked dogs and livestock straying away from herds. They preferred drinking water from rivers, and drank from lakes in seasons when water was less brackish.[8]

Sympatric carnivores

Species that were sympatric with the Caspian tiger include:

  • The Persian leopard was most likely distributed over the whole Caucasus;[35] today, it survives only in small and fragmented populations in this region.[36]
  • The jungle cat was recorded in wetlands bordering the Caspian Sea, in the Volga River delta and in valleys of the Terek, Kura, Aras, Amu Darya and Syr Darya rivers;[37][38] in southern Turkey it is known from a few coastal and inland wetlands.[39]
  • The Asiatic wildcat occurs from south-eastern Turkey, Kurdistan and the Caucasus across Central Asia to India and Mongolia in a variety of habitats.[40]

Disease

Two tigers in southwestern Tajikistan harbored 5–7 tapeworms (Taenia bubesei) in their small and large intestines.[8]

Conservation efforts

Soviet Union

In 1938, the first protected area Tigrovaya Balka (Russian: Тигровая балка, lit. 'Tiger dry creek or Tiger arroyo'), was established in Tajikistan. The name was given to this zapovednik after a tiger had attacked two Russian Army officers riding horseback along dried-up river channel known in Russian as balka. Tigrovaya Balka was apparently the last refuge of Caspian tigers in the Soviet Union, and is situated in the lower reaches of Vakhsh River between the Piandj and Kofarnihon Rivers near the border of Afghanistan. A tiger was seen there in 1958.[41]

After 1947, tigers were legally protected in the Union of Soviet Socialist Republics.[25]

Iran

In Iran, Caspian tigers had been protected since 1957, with heavy fines for shooting. In the early 1970s, biologists from the Department of Environment searched several years for Caspian tigers in the uninhabited areas of Caspian forests, but did not find any evidence of their presence.[9]

Central Asian reintroduction
Main article: Siberian Tiger Introduction Project

Stimulated by recent findings that the Siberian tiger (Amur population) is the closest relative of the Caspian tiger, albeit slightly bigger than it, discussions started as to whether the Amur tiger could be an appropriate subspecies for reintroduction into a safe place in Central Asia.[42] The Amu-Darya Delta was suggested as a potential site for such a project. A feasibility study was initiated to investigate if the area is suitable and if such an initiative would receive support from relevant decision makers. A viable tiger population of about 100 animals would require at least 5,000 km2 (1,900 sq mi) of large tracts of contiguous habitat with rich prey populations. Such habitat is not available at this stage and cannot be provided in the short term. The proposed region is therefore unsuitable for the reintroduction, at least at this stage.[25]

While the restoration of the Caspian tiger has stimulated discussions, the locations for the tiger have yet to become fully involved in the planning. But through preliminary ecological surveys it has been revealed that some small populated areas of Central Asia have preserved natural habitat suitable for tigers.[43]

In captivity
Colour-enhanced photo of the captive tiger in Berlin Zoo, 1899

Three tigers from Mazandaran were part of a collection of animals that the Persian Qajari Shah Naser al-Din used to found Tehran Zoological Garden in the 19th century.[44] A tiger from the Caucasus was housed at Berlin Zoo in the same century.[1] DNA from a tiger caught in northern Iran and housed at Moscow Zoo in the 20th century was used in the genetic test that established the Caspian tiger's close relationship with the Siberian tiger.[7]

In culture
  • During the Roman Empire, tigers and other large mammals were used in combats during gladiatorial games.[45]
  • The babr (Persian: ببر‎, tiger) features in Persian and Central Asian culture. The name "Babr Mazandaran" is sometimes given to a prominent wrestler.[10] The Persian people are apparently represented by the tiger in a Syrian mosaic from Palmyra, which commemorates the victory of Palmyrene King Odaenathus over them. The inscription conceals an earlier one that read: (Mrn), which is a title used by Odaenathus.[46] It has been suggested that it celebrates Odaenathus' victory over the Persians, the archer representing Odaenathus and the tigers the Persians; Odaenathus is about to be crowned with victory by the eagle flying above him.[47]
  • In the Fables of Pilpay, the tiger is described as furious and avid to rule over wilderness.[48]
  • In the Taurus Mountains, stone traps were used to capture leopards and tigers.[49]
gollark: I don't know digital signal processing, so my code just does it in the very naive way I thought of after failing to get a FFT to work.
gollark: That was not it. Or at least not only it.
gollark: Oh wait. It *should* be sin and not cos. Maybe that's it?
gollark: It's labelled on the top as being about 4 seconds long.
gollark: Hmm, maybe I should try rescaling it, that could work.

See also

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