Evolutionary history and distribution of African wildcat, Felis lybica in Iran

Document Type : Research Paper


1 Department of Environmental Sciences, Faculty of Natural Resources and Environment, Science and Research Branch, Islamic Azad University, Tehran, Iran

2 Department of Environment, Faculty of Natural Resources, University of Guilan, Rasht, Iran

3 Department of Environment, Faculty of Fisheries and Environmental Sciences, Gorgan University of Agricultural Sciences and Natural Resources, Gorgan, Iran



Wildcat, Felis lybica is one of the least-known cat species in Iran, in spite of the widespread distribution in the country. The molecular overview of the species using concatenated two mitochondrial markers (Cyt b, NADH5) with sampling throughout the range of the species in the country was examined. Mitochondrial DNA analysis and geographical distribution of the haplotypes indicated two well-supported subclades in the study area, comprising Asiatic wildcat, F. lybica ornata and African wildcat, F. lybica lybica (FST = 0.65; p-value < 0.001). Time-calibrated Bayesian phylogenetic analysis revealed divergence time between F. l. ornata and F. l. lybica dates to 340,000 years ago (HPD 95%: 192,000-489,000 years ago). Based on the results of dating phylogenetic tree, Central Asia is the origin area for distribution of Felis genus (Domestic Cat lineage). On the other hand, wildcat moved out of Central Asia towards Western Asia, Europe and Africa. Given estimated divergence times of less than one million years ago among wildcat subspecies, it seems that Pleistocene climatic fluctuations may have led to the diversification of this taxon. As our study does not prove hybridization between wildcat and its domestic congeners, further investigations should focus on this remarkable threatening factor.


Asadi Aghbolaghi, M, Ahmadzadeh, F, Kiabi, B & Keyghobadi, N 2019, The permanent inhabitant of the oak trees: phylogeography and genetic structure of the Persian squirrel (Sciurus anomalus). Biological Journal of the Linnean Society, 127: 197-212.
Asadi Aghbolaghi, M, Ahmadzadeh, F, Kiabi, B & Keyghobadi, N 2020, Evolutionary history of the     Persian squirrel (Sciurus anomalus): It emerged on the Eurasian continent in the Miocene. Zoologischer     Anzeiger, 287: 1-8.
Bandelt, HJ, Forster, P & Röhl, A 1999, Median-joining networks for inferring intraspecific phylogenies. Molecular Biology & Evolution, 16: 37-48.
Beaumont, M, Barratt, EM, Gottelli, D, Kitchener, AC, Danials, MJ, Pritchard, JK & Bruford, MW 2001, Genetic diversity and introgression in the Scottish wildcat. Molecular Ecology, 10: 319-336.
Bilgin, R, Karatas, A, Çoraman, E, Disotell, T & Morales, JC 2008, Regionally and climatically restricted patterns of distribution of genetic diversity in a migratory bat species Miniopterus schreibersii (Chiroptera: Vespertilionidae). BMC Evolutionary Biology, 209: 1471-2148.
Bouckaert, R, Heled, J, Kühnert, D, Vaughan, T, Wu, C-H, Xie, D, Suchard, MA, Rambaut, A & Drummond, A J 2014, BEAST 2: A Software Platform for Bayesian Evolutionary Analysis. PLoS Computational Biology, 10(4), e1003537. doi:10.1371/journal.pcbi.1003537.
Devillard, S, Jombart, Th, Léger, F, Pontier, D, Say, L, & Ruette, S 2013, How reliable are morphological and anatomical characters to distinguish European wildcats, domestic cats and their hybrids in France? Journal of Zoological Systematics and Evolutionary Research, 52: 154-162.
Driscoll, CA, Menotti-Raymond, M, Roca, AL, Hupe, K, Johnson, WE, Geffen, E, Harley, E, Delibes, M, Pontier, D, Kitchener, AC, Yamaguchi, N, O’Brien, SJ & MacDonald, D 2007, The near eastern origin of cat domestication. Science, 317: 519-523.
Driscoll, C, Yamaguchi, N, O’Brien, SJ & MacDonald, DW 2011, A suite of genetic markers useful in assessing wildcat (Felis silvestris spp.) domestic cat (Felis silvestris catus) admixture. Journal of Heredity 102 (Suppl. 1): 87-90.
Dubey, S, Cosson, J, Vohralik, V, Krystufek, B, Diker, E & Vogel, P 2007, Molecular evidence of Pleistocene bidirectional faunal exchange between Europe and the Near East: the case of the bicolored shrew (Crocidura leucodon, Soricidae). Journal of Evolutionary Biology, 20: 1799-1808.
Excoffier, L & Lischer, HEL 2010, Arlequin suite ver 3.5: a new series of programs to perform population genetics analyses under Linux and Windows. Molecular Ecology Resources, 10: 564-567.
Guggisberg, W 1975, Wild cats of the world. The University of Michigan. Taplinger Pub Co, pp. 253.
Gür, H 2013, The effects of the Late Quaternary glacial–interglacial cycles on Anatolian ground squirrels: range expansion during the glacial periods? Biological Journal of the Linnean Society, 109: 19-32.
Ghoddousi, A, kh, Hamidi, A, Ghadirian, T & Baniasadi, S 2016, The Status of wildcat in Iran: a crossroad of subspecies? Cat-News, Special Issue 10: 60-63.
Gür, H, Perktaş, U Gür, MK 2018, Do climate-driven altitudinal range shifts explain the intraspecific diversification of a narrow ranging montane mammal, Taurus ground squirrels? Mammal Research, 63: 197-211.
Hewitt, G 1999, Post-glacial re-colonization of the European biota. Biological Journal of the Linnean Society, 68: 87-112.
Hemmer, H 1978a, The evolutionary systematic of living Felidae: present status and current problems. Carnivore, 1:71-79.
Jansson, R & Dynesius, M 2002, The fate of clades in a world of recurrent climatic change: Milankovitch oscillations and evolution. Annual Review of Ecology and Systematic, 33: 741-777.
Johnson, WE & O'Brien, SJ 1997, Phylogenetic reconstruction of the felidae using 16S rRNA and NADH-5 mitochondrial genes. Journal of Molecular Evolution, 44: 98-116.
Kasapidis, P, Suchentrunk, F, Magoulas, A & Kotoulas, G 2005, The shaping of mitochondrial DNA phylogeographic patterns of the brown hare (Lepus europaeus) under the combined influence of late Pleistocene climatic fluctuations and anthropogenic translocations. Molecular Phylogenetic and Evolution, 34: 55-66.
Kilshaw, K, Johnson, P J, Kitchener, AC, & Macdonald, DW 2014, Detecting the elusive Scottish wildcat (Felis silvestris silvestris) using camera trapping. Oryx, 49: 1-9.
Kitchener, AC, Yamaguchi, N, Ward, JM, &Macdonald, DW 2005, A diagnosis for the Scottish wildcat (Felis silvestris): a tool for conservation action for a critically-endangered felid. Animal Conservation, 8: 223-237.
Kitchener, AC, Breitenmoser-Würsten, C, Eizirik, E, Gentry, A, Werdelin, L, Wilting, A, Yamaguchi, N & Johnson, WE 2017, A revised taxonomy of the Felidae. The final report of the Cat Classification Task Force of IUCN/SSC Cat Specialist Group. Cat News, Special Issue 11: 1-80.
Leigh, JW, & Bryant, D 2015, PopART: Full-feature software for haplotype network construction. Methods in Ecology and Evolution, 6: 1110-1116.
Mirzakhah, M, Naderi, S, Rezaei, HR, Fadakar, D & Naseri, M 2015, Phylogeny of gazelles in some islands of Iran based on mtDNA sequences: Species identification and implications for conservation. Caspian Journal of Environmental Sciences,13:21-30.
Mattucci, F, Oliveira, R, Lyons, LA, Alves, PC, Randi, E 2015, European wildcat populations are subdivided into five main biogeographic groups: consequences of Pleistocene climate changes or recent anthropogenic fragmentation? Ecology and Evolution, 6: 3-22.
Mousavi, M, Rezaei, HR & Naderi, S 2019, Phylogeographic analysis of Iranian wildcats (Felis lybica/Felis silvestris) as revealed by mitochondrial cytochrome b gene. Zoology in the Middle East, 65: 293-306.
Nowell, K & Jackson, P 1996, Wild cats: Status survey and conservation action plan. Gland (Switzerland): IUCN.
Nyakatura, K & Bininda-Emonds, O 2012, Updating the evolutionary history of Carnivora (Mammalia): a new species-level supertree complete with divergence time estimates. BMC Biology, 10: 1-31.
O’Brien, SJ, Johnson, W, Driscoll, C, Pontius, J, Pecon-Slattery, J & Menotti-Raymond, M 2008, State of cat genomics. PubMed Central (PMC), 24: 268-279.
O’Brien, J, Devillard, S, Say, L,Vanthomme, H, Leger, F, Ruette, S, & Pontier, D 2009, Preserving genetic integrity in a hybridizing world: are European Wildcats (Felis silvestris silvestris) in eastern France distinct from sympatric feral domestic cats? Biodiversity and Conservation, 18: 2351-2360.
Ottoni, C, Van Neer, W, De Cupere, B, Daligault, J, Guimaraes, S, Peters, J, … & Bălăşescu, A 2017, The palaeogenetics of cat dispersal in the ancient world. Nature Ecology & Evolution, 1: 139.
Pierpaoli, M, Biro, ZS, Herrmann, M, Hupe, SK, Fernandes, M, Ragni, B, Szemethy, L & Randi, E 2003, Genetic distinction of wildcat (Felis silvestris) populations in Europe, and hybridization with domestic cats in Hungary. Molecular Ecology, 12: 2585-2598.
Parmesan, C 2006, Ecological and evolutionary responses to recent climate change. Annual Review of Ecology, Evolution, and Systematic, 37: 637-669.
Posada, D 2008, JModelTest: phylogenetic model averaging. Molecular Biology Evolution, 25: 1253-1256.
Randi, E & Ragni, B 1991, Geneic variability and biochemical systematic of Domestic and Wild Cat populations (Felis silvestris: Felidae). Journal of Mammalogy, 72:79-88.
Ragni, B, & Possenti, M 1996, Variability of coat-colour and markings system in Felis silvestris. Italian Journal of Zoology, 63: 285-292.
Root, TL, Price, J, Hall, KR, Schneider, SH, Rosenweig, C & Pounds, JA 2003, Fingerprints of global warming on wild animals and plants. Nature, 421: 57-60.
Ronquist, F & Huelsenbeck, JP 2003, MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19: 1572-1574.
Senn, HV, Ghazali, M, Kaden, J, Barclay, D, Harrower, B, Campbell, RD, … & Kitchener, AC 2018, Distinguishing the victim from the threat: SNP-based methods reveal the extent of introgressive hybridization between wildcats and domestic cats in Scotland and inform future in situ and ex situ management options for species restoration. Evolutionary Applications, 12: 399-414.
Stewart, JR, Lister, AM, Barnes, I & Dalen, L 2010, Refugia revisited: individualistic responses of species in space and time. Proceedings of the Royal Society, 277: 661-671.
Stamatakis, A 2014, RAxML Version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30: 1312-1313.
Taberlet, P, Fumagalli, L, Wust-Saucy, AG & Cosson, JF 1998, Comparative phylogeography and postglacial colonization routes in Europe. Molecular Ecology, 7: 453-464.
Tamada, T, Kurose, N & Masuda, R 2005, Genetic diversity in domestic Felis catus of the Tsushima islands, based on mitochondrial DNA cytochrome b and control region nucleotide sequence. Zoological Science, 2: 627-633.
Tamura, K, Stecher, G, Peterson, D, Filipski, A & Kumar, S 2013, MEGA6: Molecular evolutionary genetics analysis version 6.0. Molecular Biology Evolution, 30: 2725-2729.
Veith, M, Schmidtler, JF, Kosuch, J, Baran, I & Seitz, A 2003, Palaeoclimatic changes explain Anatolian mountain frog evolution: a test for alternating vicariance and dispersal events. Molecular Ecology, 12: 185-199.
Wardelin, L, Yamaguchi, N, Johnson, WE & O'Brien, SJ 2010, Phylogeny and evolution of cats (Felidae). In book: Biology and conservation of wild felids, Oxford University Press, pp. 59-82.
Wozencraft, WC 2005, Order Carnivora. In Wilson, D.E. & Reeder, D.M. (eds.) Mammal Species of the World, Third Edition. The Johns Hopkins University Press, Baltimore: pp. 532-628.
Yamaguchi, N, Driscoll, CA, Kitchener, AC, Ward, JM & Macdonald, DW 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 83: 47-63.
Yamaguchi, N, Kitchener, A, Driscoll, C & Nussberger, B 2015, Felis silvestris. The IUCN red list of threatened species 2015: e.T60354712A50652361, http://dx.doi.org/10.2305/IUCN.UK. 2015-2.RLTS.T60354712 A50652361.en.
Zwijacz-Kozica, T, Wazna, A, Munoz-Fuentes, V & Tiesmeyer, A 2017, Not European wildcats, but domestic cats inhabit Tatra National Park. Polish Journal of Ecology, 65: 415-421.