Russian Journal of Theriology. Main page    

Russian Journal of Theriology. Main page
Free access to the published articles
Information about online submission, Articles format, Instructions for authors etc
Instructions for reviewers
Subscription and prices

Русскоязычный вариант сайта
Taxonomic status of the leopard, Panthera pardus (Carnivora, Felidae) in the Caucasus and adjacent areas
Khorozyan I.G., Baryshnikov G.F., Abramov A.V.
P. 41-52
Nine subspecies names have been designated for the leopard in the Middle East: Arabian or South Arabian Panthera pardus nimr (Hemprich & Ehrenberg, 1833); Anatolian or Asia Minor P. p. tulliana (Valenciennes, 1856); Caucasian P. p. ciscaucasica (Satunin, 1914); Persian or North Persian P. p. saxicolor Pocock, 1927; Sind or Baluchistan P. p. sindica Pocock, 1930; Kashmir P. p. millardi Pocock, 1930; Sinai P. p. jarvisi Pocock, 1932; Central Persian P. p. dathei Zukowsky, 1959 and South Caucasian P. p. transcaucasica Zukowsky, 1964. We have measured or retrieved data from literature on 24 characters and 3 indices of 40 leopard skulls originated from this region. We used multiple discriminant analysis to separate 7 groups from North Caucasus, South Caucasus, Turkmenistan, Iran, Turkey, Pakistan and Arabia with over 90% probabilities of correct group membership. Five scenarios of grouping were obtained which have shown the identity of the leopards from the Caucasus and Turkmenistan and their possible identity with individuals from northern Iran, closeness of leopards from southern Iran and Pakistan and from Turkey and Sinai Peninsula, and clear distinctiveness of the leopards from Arabia and Turkey from all other groups. In compliance with criteria of priority in zoological nomenclature, we suggest to retain the names P. p. ciscaucasica (=saxicolor, transcaucasica) for the Caucasus, Turkmenistan and northern Iran, P. p. tulliana for south-western Turkey, P. p. sindica (=dathei) for southern Iran and southern Pakistan and P. p. nimr for Arabian Peninsula. The subspecies P. p. millardi is probably synonymous to P. p. sindica and its status should be clarified on additional data. The taxonomic position of P. p. jarvisi should be verified by comparison with nominotypical P. p. pardus from Egypt.

DOI: 10.15298/rusjtheriol.5.1.06


  • Akin A. 1991. The status of the leopard in Turkey // 1990 International Leopard Studbook. P.7-10.
  • Alekperov A.M. 1947. [New data about the South Caucasus leopard (Panthera pardus tullianus Valenc.)] // Doklady Akademii Nauk Azerbaidzhanskoi SSR. Vol.3. No.3. P.126-128 [in Russian].
  • Anonymous. 1991. The subspecies question // Cat News. No.15. P.17-18.
  • Baryshnikov G. 2000. A new subspecies of the honey badger Mellivora capensis from Central Asia // Acta Theriologica. Vol.45. No.1. P.45-55.
  • Baryshnikov G.F. in press. [Fauna of Russia and Adjacent Territories. Mammals. Vol.1. Part 5.Family Ursidae]. Sankt-Peterburg: Nauka. [in Russian].
  • Baytop T. 1973. Neue Beobachtungen über die Verbreitung des kleinasiatischen Leoparden (Panthera pardus tulliana) in der Türkei // Bonner Zoologische Beiträge. Bd.24. Hf.3. S.183-184.
  • Birula A.A. 1912. [Materials on systematics and geographical distribution of mammals. III. Carnivora collected by N.A. Zarudny in Persia in 1896, 1898, 1900-1901, and 1903-1904] // Ezhegodnik Zoologicheskogo Muzeya Imperatorskoi Akademii Nauk. No.17. P.219-280 [in Russian].
  • Borner M. 1977. Leopards in western Turkey // Oryx. Vol.14. No.1. P.26-30.
  • Burchak-Abramovich N.O. & Jafarov R.D. 1949. [The leopard (Panthera pardus tullianus Valenc.) on Apsheron Peninsula] // Trudy Estestvenno-istoricheskogo Muzeya Akademii Nauk Azerbaidzhanskoi SSR. No.3. P.86-104 [in Russian].
  • Dal S.K. 1954. [Animal World of Armenian SSR. Vol.1. Vertebrates]. Yerevan: Izdatel'stvo Akademii Nauk Armyanskoi SSR. 415 p. [in Russian]
  • Dinnik N.Ya. 1914. [Mammals of the Caucasus. Part II. Carnivores]. Tiflis: Tipografiya K.P. Kozlovskogo. 537 p. [in Russian].
  • Gadjiev D.V. 2000. [Leopard - Panthera pardus L.] // Kerimly T. (ed.). [Animal World of Azerbaijan. T.3]. Baku: Elm. P.577-578 [in Russian].
  • Gasparyan K.M. & Agadjanyan F.S. 1974. [On the leopard in Armenia] // Biologicheskii Zhurnal Armenii. T.27. No.12. P.84-88 [in Russian].
  • Harrison D.L. & Bates P.J.J. 1991. The Mammals of Arabia. Second Edition. Sevenoaks: Harrison Zoological Museum Publication. 354 p.
  • Hatt R.T. 1959. The mammals of Iraq // Miscellaneous Publications of the Museum of Zoology, University of Michigan. No.106. P.1-113.
  • Heptner V.G. & Sludsky A.A. 1972. [Mammals of the Soviet Union. Vol.2. Part 2. Carnivores (Hyenas and Cats)]. Moskva: Vysshaya Shkola. 551 p. [in Russian]
  • ICZN. 1999. International Code of Zoological Nomenclature. Fourth Edition. London: International Trust of Zoological Nomenclature. 306 p.
  • IUCN. 2006. The 2006 IUCN Red List of Threatened Species.
  • Jafarov R.D. 1946. [The leopard (Felis pardus L.) on Apsheron] // Izvestiya Akademii Nauk Azerbaidzhanskoi SSR, Otdelenie Geologo-khimicheskikh Nauk i Nefti. T.2. No.6. P.106-107 [in Russian].
  • Khorozyan I.G. & Abramov A.V. 2005. [The spatial structure and conservation of the Persian leopard population in Armenia] // Rozhnov V.V. & Tembotova F.A. (eds.). [Mammals of the Mountainous Areas]. Moskva: KMK Scientific Press. P.199-202 [in Russian].
  • Khorozyan I., Malkhasyan A. & Asmaryan S. 2005. The Persian leopard prowls its way to survival // Endangered Species Update. Vol.22. No.2. P.51-60.
  • 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 // Zoology in the Middle East. No.26. P.41-47.
  • Kumerloeve H. 1956. Zur Verbreitung des Leoparden (Panthera pardus L.) in Anatolien // Der Zoologische Garten, Neue Folge. Bd.22. Hf.2. S.154-162.
  • Kumerloeve H. 1967. Zur Verbreitung kleinasiatischer Raub- und Huftiere sowie einiger Großnager // Säugetierkundliche Mitteilungen. Bd.15. Hf.4. S.337-409.
  • Kumerloeve H. 1976. Leoparden, Panthera pardus tulliana (Valenciennes, 1856), in Zentralanatolien // Säugetierkundliche Mitteilungen. Bd.24. Hf.1. S.46-48.
  • Larson S.E. 1997. Taxonomic re-evaluation of the jaguar // Zoo Biology. No.16. P.107-120.
  • Lukarevsky V.S. 2001. [The Leopard, Striped Hyena, and Wolf in Turkmenistan]. Moskva: Signar. 128 p. [in Russian]
  • Lukarevsky V.S. 2006. [The spotted hostage. Home for the leopard] // Panda Times. May 2006. P.8-9 [in Russian].
  • Lukarevsky V., Askerov E. & Hazaryan G. 2004. Condition of the leopard population in the Caucasus // Beiträge zur Jagd- und Wildforschung. No.29. P.305-319.
  • Lydekker R. 1899. On the leopard of the Caucasus // Proceedings of the Zoological Society of London. P.795-796.
  • Meijaard E. 2004. Biogeographic history of the Javan leopard Panthera pardus based on a craniometric analysis // Journal of Mammalogy. Vol.85. No.2. P.302-310.
  • Mendelssohn H. 1989. Felids in Israel // Cat News. No.10. P.2-4.
  • Miththapala S. 1992. Genetic and morphological variation in the leopard (Panthera pardus): a geographically widespread species // PhD dissertation. Gainesville: University of Florida.
  • Miththapala S., Seidensticker J. & O'Brien S.J. 1996. Phylogeographic subspecies recognition in leopards (Panthera pardus): molecular genetic variation // Conservation Biology. Vol.10. No.4. P.1115-1132.
  • O'Brien S.J. & Mayr E. 1991. Bureaucratic mischief: recognizing endangered species and subspecies // Science. Vol.251. No.4998. P.1187-1188.
  • Ognev S.I. 1935. [Mammals of the USSR and Adjacent Countries. T.3. Carnivores and Pinnipeds). Moskva-Leningrad: Biomedgiz. 752 p. [in Russian]
  • Ortolani A. 1999. Spots, stripes, tail tips and dark eyes: predicting the function of carnivore colour patterns using the comparative method // Biological Journal of the Linnean Society. Vol.67. No.4. P.433-476.
  • Pocock R.I. 1927. Description of two subspecies of leopards // Annals and Magazine of Natural History, Series 9. No.20. P.213-214.
  • Pocock R.I. 1932. The leopards of Africa // Proceedings of the Zoological Society of London. No.2. P.542-591.
  • Pocock R.I. 1930a. The panthers and ounces of Asia // Journal of the Bombay Natural History Society. Vol.34. No.1. P.65-82.
  • Pocock R.I. 1930b. The panthers and ounces of Asia. Part II. The panthers of Kashmir, India, and Ceylon // Journal of the Bombay Natural History Society. Vol.34. No.2. P.307-336.
  • Puzachenko A.Yu. 1993. On the taxonomic status of Felis silvestris Schreber, 1777 on the Caucasus with some comments on the variation between the European and the African wildcat // Proceedings of the Seminar on Biology and Conservation of the Wildcat (Felis silvestris), Nancy, France. Strasbourg: Council of Europe. P.75-81.
  • Quinn G.P. & Keough M.J. 2002. Experimental Design and Data Analysis for Biologists. Cambridge: Cambridge University Press. 537 p.
  • Raffel M. 2004. Persian Leopard (Panthera pardus saxicolor) EEP Annual Report 2004. Munster: Allwetter Zoo. 17 p.
  • Satunin K.A. 1905. [The mammals of Talysh and Mugan] // Izvestiya Kavkazskogo Muzeya. T.2. No.2-4. P.87-402 [in Russian].
  • Satunin K.A. 1914. [Key of the Mammals of the Russian Empire. T.1. Chiroptera, Insectivora, and Carnivora]. Tiflis: Tipografiya kantseliarii namestnika E.I.V. na Kavkaze. 184 p. [in Russian]
  • Satunin K.A. 1915. [Mammals of the Caucasus Region. T.1. Chiroptera, Insectivora, and Carnivora] // Zapiski Kavkazskogo Muzeya, Seriya A. No.1. P.1-410 [in Russian].
  • Shoemaker A.H. 1993. The Status of the Leopard, Panthera pardus, in Nature: A Country by Country Analysis. Columbia SC: Riverbanks Zoological Park. 51 p.
  • Sludsky A.A. 1973. [Distribution and abundance of wild cats in the USSR] // Trudy Instituta Zoologii Akademii Nauk Kazakhskoi SSR. No.34. P.5-106 [in Russian].
  • Thomas O. 1911. The mammals of the tenth edition of Linnaeus; an attempt to fix the types of the genera and the exact bases and localities of the species // Proceedings of the Zoological Society of London. P.120-158.
  • Uphyrkina O., Johnson W.E., Quigley H., Miquelle D., Marker L., Bush M. & O'Brien S.J. 2001. Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus // Molecular Ecology. Vol.10. No.11. P.2617-2633.
  • Valenciennes M.A. 1856. Sur une nouvelle espèce de Panthère tuée par M. Tchihatcheff à Ninfi, village situé à huit lieues est de Smyrne // Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences. No.42. P.1035-1039.
  • Werdelin L. & Olsson L. 1997. How the leopard got its spots: a phylogenetic view of the evolution of felid coat patterns // Biological Journal of the Linnean Society. Vol.63. No.3. P.383-400.
  • Yamaguchi N., Driscoll C.A., Kitchener A.C., Ward J.M. & Macdonald D.W. 2004. Craniological differentiation between European wildcats (Felis silvestris silvestris), African wildcats (F. s. lybica) and Asian wildcats (F. s. ornata): implications for their evolution and conservation // Biological Journal of the Linnean Society. Vol.83. No.1. P.47-63.
  • Zehzad B., Kiabi B.H. & Madjnoonian H. 2002. The natural areas and landscapes of Iran: an overview // Zoology in the Middle East. No.26. P.7-11.
  • Zukowsky L. 1959. Persische Panther // Der Zoologische Garten, Neue Folge. Bd.24. Hf.5-6. S.329-344.
  • Zukowsky L. 1964. Weitere Mitteilungen über Persische Panther // Der Zoologische Garten, Neue Folge. Bd.28. Hf.4. S.151-182.

Download PDF