Panthera pardus tulliana
Panthera pardus tulliana is a leopard subspecies native to the Iranian Plateau and the surrounding areas in Iran, Turkey, the Caucasus, Southern Russia, Turkmenistan and Afghanistan. Since 2016, it has been listed as Endangered on the IUCN Red List, as the population was estimated at 871 to 1,290 mature individuals, with a declining trend.
|Panthera pardus tulliana|
P. p. tulliana
|Panthera pardus tulliana|
|Distribution of P. p. tulliana, 2016|
Felis tulliana was the scientific name proposed by Achille Valenciennes in 1856, who described a skin and skull from a leopard killed near Smyrna, in western Turkey. In the 19th and 20th centuries, several naturalists described leopard zoological specimens from the Middle East:
- Felis pardus tulliana was proposed by Richard Lydekker in 1899 after examining a leopard skin from the Caucasus.
- Felis ciscaucasica was proposed by Konstantin Alekseevich Satunin in 1914, based on a leopard specimen from the Kuban region in the North Caucasus.
- Panthera pardus saxicolor was proposed by Reginald Innes Pocock in 1927, who described leopard skins from different areas of Persia, but recognized their similarity to Caucasian leopard skins. His holotype was a skin and a skull of a male leopard from Asterabad.
- P. p. sindica was proposed by Pocock in 1930 for a single skin and two skulls from the Kirthar Mountains in Balochistan. He wrote that the skin closely resemble those of P. p. saxicolor, but was distinguishable from the typical Indian leopard (P. p. fusca) in colour. It was subsumed to P. p. saxicolor based on molecular genetic analysis in 1996.
In the 19th and 20th centuries, the Anatolian leopard was considered a distinct leopard subspecies that occurred only in western Turkey. The leopard specimens from the Middle East available in zoological museum collections do not differ significantly in the sizes and shapes of skulls. Therefore, the subspecific names tulliana, ciscaucasica and saxicolor are currently considered synonyms. An analysis of leopard samples from Afghanistan revealed that they belong to P. p. saxicolor, but intergrade with the Indian leopard (P. p. fusca) in eastern Afghanistan.
In 2017, the Persian leopard population was subsumed to P. p. tulliana, which is the oldest available name for the leopard subspecies in northwestern Asia.
A phylogenetic analysis indicates that P. p. tulliana matrilineally belongs to a monophyletic group that diverged from the African (P. p. pardus) and the Arabian leopard (P. p. nimr) in the second half of the Pleistocene.DNA analysis indicates that the leopards in Iran belong to a single gene pool and form a distinct subclade.
P. p. tulliana has a grayish, slightly reddish fur with large rosettes on the flanks and back, smaller ones on the shoulder and upper legs, and spots on the head and neck. It varies in colouration; in Iran both pale and dark individuals occur. Its average body length is 158 cm (62 in), with a 192 mm (7.6 in) long skull and a 94 cm (37 in) long tail. It weighs up to 60 kg (130 lb).
Biometric data collected from 25 female and male individuals in various provinces of Iran indicate an average body length of 259 cm (102 in). A young male from northern Iran weighed 64 kg (141 lb).
Distribution and habitatEdit
The habitat of P. p. tulliana in the Greater Caucasus is subalpine meadows, temperate broadleaf and mixed forests and rugged ravines from 600 to 3,800 m (2,000 to 12,500 ft) ; and in the Lesser Caucasus and Iran rocky slopes, mountain steppes and sparse juniper forests. It avoids areas with long-duration snow cover and areas near urban development.
In northern Anatolia, zoologists found evidence of leopards in the upper forest and alpine zones of the Pontic Mountains during surveys carried out between 1993 and 2002. Its presence in the Pontic Mountains was questioned in 2016 due to a lack of evidence. A camera trap photograph obtained in Trabzon Province in the Black Sea region in September 2013 is said to show a leopard. Its preferred habitat is thought to be sparse forest areas, followed by rocky areas, agriculture and pasture areas, and riparian zones. In southeastern Turkey, its presence was documented in the Çınar district of Diyarbakır Province and in Bitlis Province. In 2018 and 2019, it was photographed on the northern slopes of Mount Cudi in Şırnak Province, and this may be a corridor for movement between Turkey and Iraq.
In the Caucasus, leopards were sighted around the Tbilisi area and in the Shida Kartli province in Georgia, where they live primarily in dense forests. Several individuals were sighted in the lowland plains of the Kakheti region in 2004. Leopard signs were also found at two localities in Tusheti, the headwaters of the Andi Koysu and Assa rivers bordering Dagestan. Between October 2000 and July 2002, tracks of 10 leopards were found in an area of 780 km2 (300 sq mi) in the rugged and cliffy terrain of Khosrov State Reserve on the southwestern slopes of the Gegham mountains. During surveys in 2013–2014, camera traps recorded leopards in 24 locations in southern Armenia, of which 14 are in the Zangezur Mountains. This trans-boundary mountain range provides important breeding habitat for leopards in the Lesser Caucasus. In March 2007 and in October 2012, an individual was photographed by a camera trap in Hirkan National Park. This protected area in southeastern Azerbaijan is in the Talysh Mountains, which are contiguous with the Alborz Mountains in Iran. During surveys in 2013–2014, camera traps recorded leopards in five locations in Hirkan National Park. The first male leopard crossing from Hirkan National Park into Iran was documented in February 2014. It was killed in the Chubar Highlands in north-western Iran's Gilan Province by a local hunter. This incident indicates that the Talysh Mountains are an important corridor for trans-boundary movement of leopards. In September 2012, the first female leopard was photographed in Zangezur National Park close to the border with Iran. During surveys in 2013–2014, camera traps recorded leopards in seven locations in Zangezur National Park, including two different females and one male. All sites are close to the border with Iran. Five cubs were documented in two sites in the Lesser Caucasus and the Talysh Mountains. Between July 2014 and June 2018, four leopards were identified in the Talysh Mountains and 11 in the trans-boundary region of Nakhchivan and southern Armenia.
Leopards have been sporadically recorded in northern Iraq. In October 2011 and January 2012, a leopard was photographed by a camera trap on Jazhna Mountain in the Zagros Mountains forest steppe in the Kurdistan Region. Between 2001 and 2014, at least nine leopards were killed by local people in this region. In 2020, a leopard was recorded in the mountains of the Sulaymaniyah Governorate in the country's northeast.
Iran is considered a stronghold of the leopard in the region. It is more abundant in the northern than in the southern part of the country, and in the 2010s was recorded in 74 of 204 protected areas. The Caspian Hyrcanian mixed forests along the Alborz Mountains are one of the most important habitats for the leopard in the country. Most leopards were recorded in habitats with temperatures of 13 to 18 °C (55 to 64 °F), maximum 20 days of ice cover per year and an annual rainfall of more than 200 mm (7.9 in). The Central Alborz Protected Area covering more than 3,500 km2 (1,400 sq mi) is one of the largest reserves in the country where leopards roam. Evidence for breeding of leopards was documented in six localities inside protected areas in the Iranian part of the Lesser Caucasus. In northeastern Iran, four leopard families with two cubs each were identified during a survey carried out from 2005 to 2008 in Sarigol National Park. A male leopard was photographed in January 2008 spraying urine on a Berberis tree; he was photographed several times until mid-February 2008 in the same area. Camera trapping surveys in summer 2016 documented the presence of 52 leopards in Sarigol, Salouk and Tandooreh National Parks. These included 10 cubs in seven families, thus highlighting that the Kopet Dag and Aladagh Mountains are important leopard refugia in the region. Leopards were also photographed in a protected area in Sefid Kuh, Kermanshah in 2020. Between September 2014 and August 2016, two radio-collared leopards moved from Iran's Kopet Dag region into Turkmenistan, revealing that the leopard population in the two countries is connected. Leopards were recorded by camera traps in the Badkhyz Nature Reserve in the country's south-west. In 2017, a young male leopard from Iran's Tandooreh National Park dispersed to and settled in Turkmenistan.  In 2018, an old male Persian leopard had moved 20 km (12 mi) from Iran to Turkmenistan. In Bamu National Park in Fars Province, surveys carried out from autumn 2007 to spring 2008 revealed seven individuals in a sampling area of 321.12 km2 (123.99 sq mi). One individual was recorded by a camera trap in Afghanistan's Bamyan Province in 2011.
In Kazakhstan, a leopard was recorded for the first time in 2000 in Jambyl Region. In 2007 and 2015, two leopards were killed in Mangystau Region farther west in the country. Between September and December 2018, camera traps recorded a leopard on a cliff in Ustyurt Nature Reserve.
P. p. tulliana was most likely distributed over the entire Caucasus, except for steppe areas. The northern foothills of the Greater Caucasus formed the northern boundary of its historic range. During surveys conducted between 2001 and 2005, no leopard was recorded in the western Greater Caucasus; it probably survived only at a few sites in the eastern part. In Armenia, people and leopards have co-existed since prehistoric times. In the mid-20th century, the leopard was relatively common in the country's mountains.
Stone traps for leopards dating to the Roman Empire still exist in the Taurus Mountains in southern Turkey. The last leopard in Syria is reported to have been killed in 1963 in the Syrian Coastal Mountain Range.P. p. tulliana was once numerous in the Aegean Region between İzmir and Antalya, with the Beşparmak Mountains considered a stronghold. Several factors contributed to the decline of the leopard population in this region between the late 1940s and mid 1970s, including deforestation, conversion of natural habitat to orchards, road construction and killing of leopards in retaliation for preying on livestock. Since surveys were not carried out in western Turkey until the mid-1980s, biologists doubted whether leopards still survived in the region. Sighting reports from the environs of Alanya in the south of the Lycian peninsula suggested that a scattered population existed between Finike, Antalya and Alanya in the early 1990s. Fresh faecal pellets found in Mount Güllük-Termessos National Park in 1992 were attributed to an Anatolian leopard. However surveys in western Turkey between 2000 and 2004 found no contemporary evidence of leopards. Extensive trophy hunting is thought to be the prime factor for the decline of the Anatolian leopard in this area. One hunter named Mantolu Hasan killed at least fifteen leopards between 1930 and 1950. It is considered locally extinct in western Turkey since the mid-1970s. No signs of the presence of leopards were detected in Termessos National Park during surveys in 2005, and local people and national park personnel were not aware of any.
The leopard population in southern Russia had been reduced to two small and isolated populations by the 1950s, and by 2007, there were fewer than 50 individuals in the region. Since 1954, leopards were thought to be extirpated in Georgia, following kills by hunters. The political and social changes caused by the breakup of the Soviet Union in 1992 caused a severe economic crisis and weakening of formerly effective protection systems; wildlife habitats were severely fragmented, leopards were persecuted and wild ungulates were hunted. In addition, inadequate baseline data and lack of monitoring programmes made it difficult to evaluate declines of mammalian prey species. In the winter of 2003, zoologists found footprints of a leopard in Vashlovani National Park in southeastern Georgia. Camera traps recorded one young male individual several times. The individual was not recorded again between 2009 and 2014. A survey in 2019 found no leopards.
Leopards also survived in northwestern Azerbaijan in the Akhar-Bakhar section of Ilisu State Reserve in the foothills of the Greater Caucasus, but in 2007 numbers were thought to be extremely low.
In Afghanistan, the leopard is thought to inhabit the central highlands of the Hindu Kush and the Wakhan corridor, but none have been photographed. The long-lasting conflict in the country badly affected both predator and prey species, so that the national population is considered to be small and severely threatened.
Behaviour and ecologyEdit
The diet of P. p. tulliana varies depending on habitat. In southern Armenia and Iran, it preys mostly on wild goat (Capra aegagrus), mouflon (Ovis gmelini), wild boar (Sus scrofa), roe deer (Capreolus capreolus), goitered gazelle (Gazella subgutturosa), Indian crested porcupine (Hystrix indica) and European hare (Lepus europaeus). It occasionally attacks livestock and herding dogs. In Iran, the presence of leopards is highly correlated with the presence of wild goat and wild sheep. An attack by a leopard on an onager (Equus hemionus) was also recorded. In Turkey, it also preys on chamois (Rupicapra rupicapra).
The mating season lasts from mid-January to mid-February. In Sarigol National Park, three females were documented in late April to May 2008 with one to two cubs each. A female with two cubs was also photographed in the Alborz Mountains.
P. p. tulliana is threatened by poaching, depletion of prey base due to poaching, human disturbance such as presence of military and training of troops in border areas, habitat loss due to deforestation, fire, agricultural expansion, overgrazing, and infrastructure development. Between 2004 and 2007, a total of 85 leopard skins were seen being offered in markets in Kabul. Leopards were known to live on the Meghri Ridge in the extreme south of Armenia, where only one individual was camera trapped between August 2006 and April 2007, but no signs of other leopards were found during track surveys conducted over an area of 296.9 km2 (114.6 sq mi). The local prey base could support 4–10 individuals. But the combined impact of poaching, disturbance caused by livestock breeding, gathering of edible plants and mushrooms, deforestation and human-induced wild fires was so high that the tolerance limits of leopards was exceeded. Only some small and isolated populations remain in the whole Caucasus. Suitable habitat is limited, and most often situated in remote border areas. Local populations depend on immigration from source populations mainly in Iran.
In Turkey, the leopard has been killed illegally in traps and through poison. Several leopards are known of have been killed since 1974 in Beypazarı, Siirt Province, Diyarbakır Province and Tunceli Provinces. Despite ongoing efforts to reforest Turkey, as of 2020[update], the country lacks a plan to reconnect fragmented forests, which may further fragment leopard populations in the region.
In Iran, primary threats are habitat disturbances, poaching and excess of livestock in leopard habitats. Leopards are unlikely to persist outside of protected areas. Droughts in wide areas of leopard habitats affected the main prey species such as wild goat and wild sheep. An assessment of leopard mortality in Iran revealed that 71 leopards were killed between 2007 and 2011 in 18 provinces; 70% were hunted or poisoned illegally, and 18% died in road accidents. Between 2000 and 2015, 147 leopards were killed in the country. More than 60% of them died due to poaching, through poisonous bait, and were shot by rangers, trophy hunters and the military. About 26% of them died in road accidents. More males than females were killed.
In the 1980s, anti-personnel mines were deployed along the northern part of the Iran-Iraq border to deter people from entering the area. Leopards roaming the area are safe from poachers and industrial development, but at least two individuals are known to have stepped on mines and been killed. The main threat in northern Iraq is deforestation, which in the early 2020s is being worsened by an economic crisis.
|Russian North Caucasus||2008||fewer than 10|
|Georgia||2008||fewer than 5|
|Turkey||2005||fewer than 5|
Panthera pardus is listed in CITES Appendix I. It is listed as a strictly protected species in Appendix II of the Berne Convention on the Conservation of European Wildlife and Natural Habitats. In Azerbaijan, the leopard has been protected by law since 1969; in Armenia and in the Soviet Union, it was protected by law in 1972; the Caucasus leopard population was listed in Russia's Red Data Book under Category I as threatened with extinction. It has been protected by law in Iran since 1999. In 2001, hunting leopards was banned in Nakhchivan Autonomous Republic, and anti-poaching activities were regularly conducted in southern Armenia since 2003. Since 2005, seven protected areas have been established in the Lesser Caucasus covering an area of 1,940 km2 (750 sq mi), and three in the Talysh Mountains with an area of 449 km2 (173 sq mi). The total protected area in the country now amounts to 4,245 km2 (1,639 sq mi). In Georgia's national Red Data Book, the leopard has been listed as Critically Endangered since 2006. Penalties for killing leopards were adopted and increased several times in Armenia and Azerbaijan. In Afghanistan, it was included in the country's Protected Species List in 2009. In Kazakhstan, hunting was made a criminal offence in 2021.
In 2001, a 5-year leopard conservation project was initiated in the Caucasus, which supported systematic surveys in the region, the planning of new and enlargement of existing protected areas, training of border guards and school education campaigns in Armenia and Azerbaijan; an anti-poaching unit was set up in Armenia. In 2005, the Armenian Ministry of Environment approved a conservation plan with the leopard as an umbrella species, and a strategy for leopard conservation in the Caucasus in 2008. In Iran, a leopard conservation and management action plan was endorsed in 2016. As of 2019[update], Nature Iraq is mapping potential habitat near the border with Iran as the first stage of a conservation project.
In 2009, the Leopard Breeding and Rehabilitation Centre in Russia's Sochi National Park received two leopards from Turkmenistan as part of a leopard breeding and reintroduction programme. Since then, leopards have been brought from various zoos. Their offspring were released into the wild in 2016 and 2018, including three males and one female into the Caucasus Biosphere Reserve, and one male and one female into Alaniya National Park in North Ossetia–Alania.
As of December 2011, there were 112 captive Persian leopards in zoos worldwide, comprising 48 male, 50 female and five unsexed individuals less than 12 months of age within the European Endangered Species Programme.[needs update]
In 2015, representations of the Anatolian leopard were found in the ancient city of Thyatira in Anatolia that date from the Neolithic period to the end of the 6th century BC. The leopard is depicted on statues, potteries, ivory works and coins associated with the Lydian culture. Several pieces were found in areas that were used for worship.
The Natural History Museum of the Aegean on Samos Island in Greece exhibits a stuffed animal said to have been killed on the island in 1862, labelled Kaplani, Greek: Καπλάνι, meaning leopard. Its size, shape and coat colour is rather unnatural for a leopard, but may have been altered in the process of taxidermy. It is possible that an Anatolian leopard reached the island by swimming across the 1.7 km (1.1 mi) wide channel from the Turkish coast. The story of the leopard and the exhibit inspired Greek author Alki Zei to write a novel for children in Greek: Το καπλάνι της βιτρίνας, translated as Wildcat Under Glass.
- Stein, A.B.; Athreya, V.; Gerngross, P.; Balme, G.; Henschel, P.; Karanth, U.; Miquelle, D.; Rostro-Garcia, S.; Kamler, J.F.; Laguardia, A.; Khorozyan, I. & Ghoddousi, A. (2020). "Panthera pardus (amended version of 2019 assessment)". IUCN Red List of Threatened Species. 2020: e.T15954A163991139.
- Kitchener, A. C.; Breitenmoser-Würsten, C.; Eizirik, E.; Gentry, A.; Werdelin, L.; Wilting, A.; Yamaguchi, N.; Abramov, A. V.; Christiansen, P.; Driscoll, C.; Duckworth, J. W.; Johnson, W.; Luo, S.-J.; Meijaard, E.; O’Donoghue, P.; Sanderson, J.; Seymour, K.; Bruford, M.; Groves, C.; Hoffmann, M.; Nowell, K.; Timmons, Z.; Tobe, S. (2017). "A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group" (PDF). Cat News (Special Issue 11): 73–75.
- Pocock, R. I. (1930). "The Panthers and Ounces of Asia". Journal of the Bombay Natural History Society. 34 (1): 65–82.
- Ullrich, B. & Riffel, M. (1993). "New evidence for the occurrence of the Anatolian Leopard, Panthera pardus tulliana (Valenciennes, 1856), in Western Turkey". Zoology in the Middle East. 8 (1): 5–14. doi:10.1080/09397140.1993.10637631.
- Valenciennes, A. (1856). "Sur une nouvelles espèce de Panthère tué par M. Tchihatcheff à Ninfi, village situé à huit lieues est de Smyrne" [About a new Panther species that was killed by Mr. Tchihatcheff at Ninfi, a village located eight leagues from Smyrna]. Comptes rendus hebdomadaires des séances de l'Académie des sciences. 42: 1035–1039.
- Lydekker, R. (1899). "On the Leopard of the Caucasus". Proceedings of the Zoological Society of London (June): 795–796.
- Satunin, K. A. (1914). "Leopardus pardus ciscaucasicus, Satunin". Conspectus Mammalium Imperii Rossici I. Tiflis: Tipografiâ Kancelârii Namestnika E. I. V. na Kavkaz. pp. 159–160.
- Satunin, K. A. (1914). Key of the Mammals of the Russian Empire (in Russian). 1: Chiroptera, Insectivora and Carnivora. Tiflis: Tipografīi︠a︡ Kant︠s︡eli︠a︡rīi nami︠e︡stnika E.I.V. na Kavkazi︠e︡.
- Pocock, R. I. (1927). "Description of two subspecies of leopards". Annals and Magazine of Natural History. Series 9. 20 (116): 213–214. doi:10.1080/00222932708655586.
- Miththapala, S.; Seidensticker, J. & O'Brien, S. J. (1996). "Phylogeographic Subspecies Recognition in Leopards (P. pardus): Molecular Genetic Variation". Conservation Biology. 10 (4): 1115–1132. doi:10.1046/j.1523-1739.1996.10041115.x.
- Uphyrkina, O.; Johnson, E.W.; Quigley, H.; Miquelle, D.; Marker, L.; Bush, M. & O'Brien, S. J. (2001). "Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus" (PDF). Molecular Ecology. 10 (11): 2617–2633. doi:10.1046/j.0962-1083.2001.01350.x. PMID 11883877. S2CID 304770.
- Khorozyan, I. G.; Gennady, F.; Baryshnikov, G. F. & Abramov, A. V. (2006). "Taxonomic status of the leopard, Panthera pardus (Carnivora, Felidae) in the Caucasus and adjacent areas". Russian Journal of Theriology. 5 (1): 41–52. doi:10.15298/rusjtheriol.05.1.06.
- Khorozyan, I. (2014). "Morphological variation and sexual dimorphism of the common leopard (Panthera pardus) in the Middle East and their implications for species taxonomy and conservation". Mammalian Biology-Zeitschrift für Säugetierkunde. 79 (6): 398–405. doi:10.1016/j.mambio.2014.07.004.
- Manati, A. R. (2012). "Subspecies question of Persian Leopards clarified". Der Zoologische Garten. 81 (1): 1–13. doi:10.1016/j.zoolgart.2012.02.001.
- Farhadina, M. S.; Farahmand, H.; Gavashelishvili, A.; Kaboli, M.; Karami, M.; Khalili, B. & Montazamy, Sh. (2015). "Molecular and craniological analysis of leopard, Panthera pardus (Carnivora: Felidae) in Iran: support for a monophyletic clade in western Asia". Biological Journal of the Linnean Society. 114 (4): 721–736. doi:10.1111/bij.12473.
- Farhadinia, M.S.; Ashrafzadeh, M. R.; Senn, H.; Ashrafi, S.; Farahmand, H.; Ghazali, M.; Hunter, L. T. B. & MacDonald, D. W. (2020). "Is there low maternal genetic variation in West Asian populations of leopard?". Mammal Research. 65 (4): 701–708. doi:10.1007/s13364-020-00510-1. S2CID 219542899.
- Kiabi, B.H.; Dareshouri, B.F.; Ghaemi, R.A. & Jahanshahi, M. (2002). "Population status of the Persian leopard (Panthera pardus saxicolor Pocock, 1927) in Iran" (PDF). Zoology in the Middle East. 26: 41–47. doi:10.1080/09397140.2002.10637920. S2CID 53603392.
- Lukarevsky, V.; Malkhasyan, A. & Askerov, E. (2007). "Biology and ecology of the leopard in the Caucasus". Cat News (Special Issue 2): 4–8.
- Sanei, A. (2007). Analysis of leopard (Panthera pardus) status in Iran. Tehran: Sepehr Publication Center.
- Lukarevsky, V.; Akkiev, M.; Askerov, E.; Agili, A.; Can, E.; Gurielidze, Z.; Kudaktin, A.; Malkhasyan, A. & Yarovenko, Y. (2007). "Status of the Leopard in the Caucasus" (PDF). Cat News (Special Issue 2): 15–21.
- Gavashelishvili, A.; Lukarevskiy, V. (2008). "Modelling the habitat requirements of leopard Panthera pardus in west and central Asia". Journal of Applied Ecology. 45 (2): 579–588. doi:10.1111/j.1365-2664.2007.01432.x.
- Baskaya, S. & Bilgili, E. (2004). "Does the leopard Panthera pardus still exist in the Eastern Karadeniz Mountains of Turkey ?". Oryx. 38 (2): 228–232. doi:10.1017/S0030605304000407.
- Spassov, N.; Ignatov, A. & Acosta-Pankov, I. (2016). "On the status of the leopard in Turkey, again". Cat News (64): 18–21.
- Sarı, A.; Gündoğdu, E.; Başkaya, Ş. & Arpacık, A. (2020). "Habitat preference by the Anatolian leopard (Panthera pardus tulliana Valenciennes, 1856) in North-eastern Anatolia, Turkey". Belgian Journal of Zoology. 150: 153–168. doi:10.26496/bjz.2020.78.
- Avgan, B.; Raza, H.; Barzani, M. & Breitenmoser, U. (2016). "Do recent leopard Panthera pardus records from northern Iraq and south-eastern Turkey reveal an unknown population nucleus in the region?". Zoology in the Middle East. 62 (2): 95–104. doi:10.1080/09397140.2016.1173904. S2CID 88004181.
- Toyran, K. (2018). "Noteworthy record of Panthera pardus in Turkey (Carnivora: Felidae)". Fresenius Environmental Bulletin. 27 (11): 7348–7353.
- Karataş, A.; Bulut, Ş. & Akbaba, B. (2021). "Camera trap records confirm the survival of the Leopard (Panthera pardus L., 1758) in eastern Turkey (Mammalia: Felidae)". Zoology in the Middle East. 67 (3): 198–205. doi:10.1080/09397140.2021.1924419. S2CID 235564429.
- Butkhuzi, L. (2004). "Breaking news: leopard in Georgia". Caucasus Environment (2): 49–51.
- WWF (2007). Strategy for the Conservation of the Leopard in the Caucasus Ecoregion. Strategic Planning Workshop on Leopard Conservation in the Caucasus, 30 May – 1 June 2007 (PDF) (Report). Tbilisi, Georgia.
- Khorozyan, I. & Malkhasyan, A. (2002). "Ecology of the leopard (Panthera pardus) in Khosrov Reserve, Armenia: implications for conservation" (PDF). Scientific Reports of the Zoological Society "La Torbiera" (6): 1–41.
- Khorozyan, I. (2003). "Habitat preferences by the Persian Leopard (Panthera pardus saxicolor Pocock, 1927) in Armenia". Zoology in the Middle East. 30: 25–36. doi:10.1080/09397140.2003.10637984. S2CID 84530055.
- Askerov, E.; Talibov, T.; Manvelyan, K.; Zazanashvili, N.; Malkhasyan, A.; Fatullayev, P. & Heidelberg, A. (2015). "South-Eastern Lesser Caucasus: the most important landscape for conserving the leopard (Panthera pardus) in the Caucasus region (Mammalia: Felidae)". Zoology in the Middle East. 61 (2): 95–101. doi:10.1080/09397140.2015.1035003. S2CID 84687390.
- Farhadinia, M.S.; Ahmadi, M.; Sharbafi, E.; Khosravi, S.; Alinezhad, H. & Macdonald, D.W. (2015). "Leveraging trans-boundary conservation partnerships: Persistence of Persian leopard (Panthera pardus saxicolor) in the Iranian Caucasus" (PDF). Biological Conservation. 191: 770–778. doi:10.1016/j.biocon.2015.08.027.
- Бабаева, З. (2007). "Представителю Бакинского офиса Всемирного Фонда охраны дикой природы удалось впервые сфотографировать в Азербайджане живого леопарда" [According to the Representative of the WWF Baku Office it was possible for the first time to photograph a leopard in Azerbaijan]. Day (in Russian).
- Исабалаева, И. (2012). "В Гирканском национальном парке Азербайджана обнаружен еще один кавказский леопард" [One Caucasian leopard found again in Azerbaijan's Hirkan National Park]. Novosti (in Russian). Archived from the original on 2 December 2012.
- Maharramova, E.; Moqanaki, E.M.; Askerov, E.; Faezi, S.; Alinezhad, H.; Mousavi, M. & Zazanashvili, N. (2018). "Transboundary leopard movement between Azerbaijan and Iran in the Southern Caucasus" (PDF). Cat News (67): 8–10.
- Avgan, B.; Ismayilov, A.; Fatullayev, P.; Huseynali, T. T.; Askerov, E. & Breitenmoser, U. (2012). "First hard evidence of leopard in Nakhchivan". Cat News (57): 33.
- Breitenmoser, U.; Askerov, E.; Soofi, M.; Breitenmoser-Würsten, C.; Heidelberg, A.; Manvelyan, K. & Zazanashvili, N. (2017). "Transboundary leopard conservation in the Lesser Caucasus and the Alborz Range". Cat News (65): 24–25.
- Askerov, E.; Talibov, T.; Manvelyan, K.; Zazanashvili, N.; Fatullayev, P. & Malkhasyan, A. (2019). "Leopard (Panthera pardus) reoccupying its historic range in the South Caucasus: a first evidence (Mammalia: Felidae)". Zoology in the Middle East. 65 (1): 88–90. doi:10.1080/09397140.2018.1552349. S2CID 91257820.
- Al-Sheikhly, O. F. (2012). "The hunting of endangered mammals in Iraq". Wildlife Middle East (6): 2–3.
- Raza, H. A.; Ahmad, S. A.; Hassan, N. A.; Qadir, K. A. M. & Ali, L. (2012). "First photographic record of the Persian leopard in Kurdistan, northern Iraq". Cat News (56): 34–35.
- Ahmed, S.H. & Majeed, S.I. (2020). "Monitoring of the wild mammal fauna in Bamo Mountain in northern Iraq (Kurdistan) for the first time using camera trap method and raising awareness for its conservation". Bulletin of the Iraq Natural History Museum. 16 (2): 161–172. doi:10.26842/binhm.7.2020.16.2.0161.
- Sanei, A.; Mousavi, M.; Kiabi, B.H.; Masoud, M.R.; Gord Mardi, E.; Mohamadi, H.; Shakiba, M.; Baran Zehi, A.; Teimouri, M. & Raesi, T. (2016). "Status assessment of the Persian leopard in Iran". Cat News (Special Issue 10): 43–50.
- Sanei, A.; Zakaria, M. (2011). "Distribution pattern of the Persian leopard ("Panthera pardus saxicolor") in Iran". Asia Life Sciences (Supplement 7): 7–18.
- Farhadinia, M.; Nezami, B.; Mahdavi, A. & Kaveh, H. (2007). "Photos of Persian Leopard in Alborz Mountains, Iran" (PDF). Cat News. 46: 34–35.
- Farhadinia, M.; Mahdavi, A. & Hosseini-Zavarei, F. (2009). "Reproductive ecology of Persian leopard, Panthera pardus saxicolor, in Sarigol National Park, northeastern Iran" (PDF). Zoology in the Middle East. 48: 13–16. doi:10.1080/09397140.2009.10638361. S2CID 83357675.
- Farhadinia, M.S.; McClintock, B.T.; Johnson, P.J.; Behnoud, P.; Hobeali, K.; Moghadas, P.; Hunter, L.T. & Macdonald, D.W. (2019). "A paradox of local abundance amidst regional rarity: the value of montane refugia for Persian leopard conservation". Scientific Reports. 9 (1): 14622. Bibcode:2019NatSR...914622F. doi:10.1038/s41598-019-50605-2. PMC 6788991. PMID 31604967.
- "Hopes rise for Persian leopard survival in western Iran". Mehr News Agency. 6 February 2021. Retrieved 9 April 2021.
- Farhadinia, M.; Memarian, I.; Hobeali, K.; Shahrdari, A.; Ekrami, B.; Kaandorp, J. & Macdonald, D.W. (2017). "GPS collars reveal transboundary movements by Persian leopards in Iran". Cat News (65): 28–30.
- Kaczensky, P. & Linnell, J. D. C. (2014). Rapid assessment of the mammalian community of the Badhyz Ecosystem, Turkmenistan, October 2014. NINA Report 1148. Trondheim: Norwegian Institute for Nature Research.
- Farhadinia, M.S.; Johnson, P.J.; Macdonald, D.W. & Hunter, L.T.B. (2018). "Anchoring and adjusting amidst humans: Ranging behavior of Persian leopards along the Iran-Turkmenistan borderland". PLOS ONE. 13 (5): e0196602. Bibcode:2018PLoSO..1396602F. doi:10.1371/journal.pone.0196602. PMC 5931651. PMID 29719005.
- IFP Editorial Staff (2018). "Persian Leopard Borzou Migrates to Turkmenistan". Iran Front Page.
- Ghoddousi, A.; Hamidi, A. Kh.; Ghadirian, T.; Ashayeri, D.; Khorozyan, I. (2010). "The status of the Endangered Persian leopard Panthera pardus saxicolor in Bamu National Park, Iran". Oryx. 44 (4): 551–557. doi:10.1017/S0030605310000827.
- Moheb, Z. & Bradfield, D. (2014). "Status of the common leopard in Afghanistan". Cat News (61): 15–16.
- Shakula, V. (2004). "First record of leopard (Panthera pardus) in Kazakhstan". Cat News (41): 11–12.
- Pestov, M. V.; Nurmukhambetov, Z. E.; Mukhashov, A. T.; Terentyev, V. A. & Rosen, T. (2019). "First camera trap record of Persian leopard in Ustyurt State Nature Reserve, Kazakhstan". Cat News (69): 14–16.
- Khorozyan, I. (2003). "The Persian leopard in Armenia: research and conservation". Proceedings of Regional Scientific Conference "Wildlife Research and Conservation in South Caucasus", 7–8 October 2003, Yerevan, Armenia. Yerevan. pp. 161–163.
- Sekercioglu, C.; Anderson, S.; Akcay, E.; Bilgin, R.; Can, O. & Semiz, G. (2011). "Turkey's Globally Important Biodiversity In Crisis" (PDF). Biological Conservation. 144 (12): 2752–2769. doi:10.1016/j.biocon.2011.06.025.
- Masseti, M. (2009). "Carnivores of Syria". In Neubert, E.; Amr, Z.; Taiti, S.; Gümüs, B. (eds.). Animal Biodiversity in the Middle East. Proceedings of the First Middle Eastern Biodiversity Congress, Aqaba, Jordan, 20–23 October 2008. ZooKeys. ZooKeys 31. pp. 229–252. doi:10.3897/zookeys.31.170.
- Borner M. (1977). "Leopards in western Turkey". Oryx. 14 (1): 26–30. doi:10.1017/S0030605300014769.
- Can, O. E. (2004). Status, conservation and management of large carnivores in Turkey (Report). Strasbourg: Convention on the Conservation of European Wildlife and Natural Habitats. Standing Committee, 24th meeting, 29 November–3 December 2004.
- Giannatos, G.; Albayrak, T. & Erdogan, A. (2006). "Status of the Caracal in Protected Areas in Southwestern Turkey". Cat News (45): 23–24.
- Khorozyan, I. G.; Gennady, F.; Baryshnikov, G. F. & Abramov, A. V. (2006). "Taxonomic status of the leopard, Panthera pardus (Carnivora, Felidae) in the Caucasus and adjacent areas" (PDF). Russian Journal of Theriology. 5 (1): 41–52. doi:10.15298/rusjtheriol.05.1.06.
- Mallon, D.; Weinberg, P. & Kopaliani, N. (2007). "Status of the prey species of the Leopard in the Caucasus" (PDF). Cat News (Special Issue 2): 22–27.
- Antelava, N. (2004). "Lone leopard spotted in Georgia". BBC News, 25 May 2004.
- Zazanashvili, N.; Manvelyan, K.; Askerov, E. & Heidelberg, A. (2020). Leopard in the South Caucasus (PDF). Tbilisi, Georgia: WWF-Caucasus.
- Habibi, K. (2004). Mammals of Afghanistan. Coimbatore: Zoo Outreach Organization.
- Farhadinia, M.S.; Nezami, B.; Hosseini-Zavarei, F. & Valizadeh, M. (2009). "Persistence of Persian leopard in a buffer habitat in northeastern Iran" (PDF). Cat News (51): 34–36.
- Khorozyan, I.; Malkhasyan, A. & Asmaryan, S. (2005). "The Persian leopard prowls its way to survival" (PDF). Endangered Species Update. 22 (2): 51–60. Archived from the original (PDF) on 3 July 2013.
- Lukarevsky V.; Malkhasyan, A. & Askerov, E. (2007). "Biology and ecology of the leopard in the Caucasus". Cat News (Special Issue 2): 9–15.
- Sanei, A.; Zakaria, M. & Hermidas, S. (2011). "Prey composition in the Persian leopard distribution range in Iran". Asia Life Sciences. Supplement 7: 19–30. Archived from the original on 27 December 2013. Retrieved 27 July 2012.
- Manati, A. R. (2009). "The trade in Leopard and Snow Leopard skins in Afghanistan". TRAFFIC Bulletin. 22 (2): 57–58.
- Khorozyan, I.; Malkhazyan, A. G. & Abramov, A. (2008). "Presence – absence surveys of prey and their use in predicting leopard (Panthera pardus) densities: a case study from Armenia" (PDF). Integrative Zoology. 2008 (3): 322–332. doi:10.1111/j.1749-4877.2008.00111.x. PMID 21396082. Archived from the original (PDF) on 31 July 2013. Retrieved 18 November 2012.
- Farhadinia, M.S.; Rostro-García, S.; Feng, L.; Kamler, J.F.; Spalton, A.; Shevtsova, E.; Khorozyan, I.; Al-Duais, M.; Ge, J. & Macdonald, D.W. (2020). "Big cats in borderlands: challenges and implications for transboundary conservation of Asian leopards". Oryx. 55 (3): 452–460. doi:10.1017/S0030605319000693.
- Haber7. "Bilim adamları leopar bilgisini gizli tutuyordu" [Scientists keep leopard information secret]. Haber7 (in Turkish). Retrieved 3 March 2021.
- Ertüzün, M. (2006). "The last Anatolian Panther". Mehmet Ertüzün.
- Hürriyet Daily News (2013). Shepherd kills first Anatolian leopard sighted in Turkey for years Hürriyet Daily News, 3 November 2013.
- HDN (2013). "Anatolian Leopard Sighted in Turkey for First Time in Years After Being Killed by Shepherd". Good Morning Turkey. Archived from the original on 11 November 2013.
- HDN (2013). "Mystery Anatolian Leopard Seen, Shot Before: Forensics". GoodMorningTurkey.com. Archived from the original on 11 November 2013.
- "Tunceli'de ölü Anadolu Parsı bulundu" [Dead Anatolian leopard found in Tunceli]. İhlas Haber Ajansı (in Turkish). Retrieved 3 March 2021.
- "Nesli tükenmekte olan 'Anadolu Parsı' ölü bulundu" [Endangered 'Anatolian Leopard' found dead]. www.cumhuriyet.com.tr (in Turkish). Retrieved 3 March 2021.
- Gülçi̇n, D. & Yilmaz, T. (2020). "Evaluation of forest fragmentation with particular reference to landscape-based ecological assessment and wildlife conservation". Turkish Journal of Forestry. 21 (1): 84–93. doi:10.18182/tjf.654954.
- Sanei, A. & Zakaria, M. (2009). Primary threats to Persian leopard (Panthera pardus saxicolor) in the Islamic Republic of Iran. Proceedings of the 8th International Annual Symposium on Sustainability Science and Management. 3–4 May 2009. Diterbitkan Oleh, Terengganu, Malaysia.
- Sanei, A. & Zakaria, M. (2011). "Survival of the Persian leopard (Panthera pardus saxicolor) in Iran: Primary threats and human-leopard conflicts". Asia Life Sciences. Supplement 7: 31–39. Archived from the original on 19 October 2013. Retrieved 27 July 2012.
- Sanei, A.; Mousavi, M.; Mousivand, M. & Zakaria, M. (2012). "Assessment of the Persian leopard mortality rate in Iran". Proceedings of UMT 11th International Annual Symposium on Sustainability Science and Management: 1458–1462.
- Naderi, M.; Farashi, A. & Erdi, M.A. (2018). "Persian leopard's (Panthera pardus saxicolor) unnatural mortality factors analysis in Iran". PLOS ONE. 13 (4): e0195387. Bibcode:2018PLoSO..1395387N. doi:10.1371/journal.pone.0195387. PMC 5918793. PMID 29694391.
- Schwartzstein, P. (2014). "For Leopards in Iran and Iraq, Land Mines Are a Surprising Refuge". National Geographic. Retrieved 18 March 2015.
- Crisp, W. (2020). "Last chance for the Persian leopard: the fight to save Iraqi Kurdistan's forests". The Guardian. Retrieved 11 April 2021.
- Khorozyan, I. (2008). "Panthera pardus ssp. saxicolor". IUCN Red List of Threatened Species. 2008. Archived from the original on 19 October 2009.old-form url
- Council of Europe (2002). Convention on the Conservation of European Wildlife and Natural Habitats, Appendix II. European Treaty Series No. 104. Strasbourg: Council of Europe.
- Zazanashvili, N.; Mörschel, F.; Askerov, E.; Manvelyan, K.; Krever, V.; Farvar, M. T. & Kalem, S. (2007). "Conservation of the Leopard in the Caucasus". Cat News (Special Issue 2): 4–8.
- "Kazakhstan bolsters protections for rare animal species | Eurasianet". eurasianet.org. 2021. Retrieved 9 April 2021.
- Khorozyan, I. & Malkhasyan, A., eds. (2008). National Action Plan for leopard conservation in Armenia. Yerevan: WWF Armenia Branch.
- Sanei, A. (2016). Persian Leopard National Conservation and Management Action Plan in Iran. Department of Environment of Iran. ISBN 978-600-04-4354-2.
- Sanei A.; Teimouri A.; Ahmadi Fard, G.; Asgarian, H.R. & Alikhani M. (2020). "Introduction to the Persian Leopard National Conservation and Management Action Plan in Iran". In Sanei, A. (ed.). Research and Management Practices for Conservation of the Persian Leopard in Iran. pp. 175–187. doi:10.1007/978-3-030-28003-1_8. ISBN 978-3-030-28001-7. S2CID 228086620.
- "Persian Leopard Conservation". Nature Iraq. Retrieved 5 November 2019.
- WWF (2009). "Flying Turkmen leopards to bring species back to Caucasus". WWF Russia.
- Druzhinin, A. (2010). "Iranian leopards make themselves at home in Russia's Sochi". RIA Novosti.
- "Two Leopards from Sweden arrived to Russia". WWF Russia. 2020. Retrieved 9 April 2021.
- Rozhnov, V.V.; Pshegusov, R.H.; Hernandez-Blanco, J.A.; Chistopolova, M. D.; Pkhitikov, A. B.; Trepet, S. A.; Dronova, N. A.; Naidenko, S. V. & Yachmennikova, A. A. (2020). "MaxEnt Modeling for Predicting Suitable Habitats in the North Caucasus (Russian Part) for Persian Leopard (P. p. ciscaucasica) Based on GPS Data from Collared and Released Animals". Izvestiya, Atmospheric and Oceanic Physics. 56 (9): 1090–1106. Bibcode:2020IzAOP..56.1090R. doi:10.1134/S0001433820090212. S2CID 232052406.
- International Species Information System (2011). "ISIS Species Holdings: Panthera pardus saxicolor, December 2011".
- Erön, A. (2020). "Thyateira/Hastane Höyüğü'nde Bulunan Anadolu Parsı Heykel Parçası Üzerine Düşünceler" [Thoughts about the Anatolian Leopard Statue Piece from Thyateira/Hastane Mound]. Arkhaia Anatolika. The Journal of Anatolian Archaeological Studies. 3: 82–97. doi:10.32949/Arkhaia.2020.16.
- Κρητικίδης, Ε. Ι. (1869). Τοπογραφία Αρχαία και Σημερινή της Σάμου [Ancient and Modern Topography of Samos]. Εν Ερμουπόλει: Ρενιέρη Πρίντεζη.
- Masseti, M. (2010). "Homeless mammals from the Ionian and Aegean islands". Bonn Zoological Bulletin. 57 (2): 367–373.
- Zei, A. (1963). Το Καπλ́ανι τ́ης βιτρ́ινας μυθιστ́ορημα. Athīna: Κ́ερδος.
- Zei, A. (1968). Wildcat under Glass (First ed.). New York: Holt, Rinehart & Winston.
|Wikispecies has information related to Panthera pardus tulliana.|
|Wikimedia Commons has media related to Panthera pardus tulliana.|
- "Persian leopard (Panthera pardus tulliana)". IUCN/SSC Cat Specialist Group.
- "Caucasian Leopard – The King of the Armenian mountains". WWF-Armenia.
- "ICS study confirms transboundary leopard movements between Azerbaijan and Iran in the Caucasus". Iranian Cheetah Society. 2018.
- Leopards .:. wild-cat.org — Information about research and conservation of leopards in Asia
- "Research, Conservation and Management of Asian leopard subspecies". Asian Leopard Specialist Society. 2011.
- "Pair of Persian leopards spotted for the first time in Pakistan". ARY News. 20 May 2021. Retrieved 24 May 2021.