Available online at www.sciencedirect.com Genomics 91 (2008) 12–21 www.elsevier.com/locate/ygeno The ascent of cat breeds: Genetic evaluations of breeds and worldwide random-bred populations Monika J. Lipinski a, Lutz Froenicke a, Kathleen C. Baysac a, Nicholas C. Billings a, Christian M. Leutenegger b, Alon M. Levy c, Maria Longeri d, Tirri Niini e, Haydar Ozpinar f, ⁎ Margaret R. Slater g, Niels C. Pedersen b, Leslie A. Lyons a, a Department of Population Health and Reproduction, School of Veterinary Medicine, University of California at Davis, Davis, CA 95616, USA b Department of Veterinary Medicine and Epidemiology, School of Veterinary Medicine, University of California at Davis, Davis, CA 95616, USA c Department of Molecular Genetics, Weizmann Institute of Science, Rehovot 76100, Israel d Istituto di Zootecnica, Faculty of Veterinary Medicine, University of Milan, Milan, Italy e Oy Triniini Company, P.O. Box 36, FIN-00501 Helsinki, Finland f Department of Genetics and Bioengineering, Faculty of Engineering, University of Yeditepe, 34755 Kayisdagi-Istanbul, Turkey g Department of Veterinary Integrative Biosciences, College of Veterinary Medicine, Texas A&M University, College Station, TX 77843-4458, USA Received 11 September 2007; accepted 9 October 2007 Available online 3 December 2007 Abstract The diaspora of the modern cat was traced with microsatellite markers from the presumed site of domestication to distant regions of the world. Genetic data were derived from over 1100 individuals, representing 17 random-bred populations from five continents and 22 breeds. The Mediterranean was reconfirmed to be the probable site of domestication. Genetic diversity has remained broad throughout the world, with distinct genetic clustering in the Mediterranean basin, Europe/America, Asia and Africa. However, Asian cats appeared to have separated early and expanded in relative isolation. Most breeds were derived from indigenous cats of their purported regions of origin. However, the Persian and Japanese bobtail were more aligned with European/American than with Mediterranean basin or Asian clusters. Three recently derived breeds were not distinct from their parental breeds of origin. Pure breeding was associated with a loss of genetic diversity; however, this loss did not correlate with breed popularity or age. © 2007 Elsevier Inc. All rights reserved. Keywords: Breeds; Cat; Diversity; Domestication; Genetic; Felis; Origins; Phylogeography; Phylogenetics; Structure The domestic cat (Felis catus, Linneas 1758) is prolific and possible by the domestication of certain wild grains and grasses cosmopolitan, occupying most habitable corners of the world. [4]. The human/cat relationship was beneficial for the control of Archeological evidence suggests the domestication of the cat crop-destroying rodents, which had also joined their evolution- occurred in the Near East approximately 9000–10,000 years ary fate to human civilization. Although many cats eventually ago [1]. However, the initiation of domestication probably became pets, the modern cat is not fully domesticated in the began thousands of years earlier as humans and ancestral cats classic sense. Modern cats remain self-sufficient if need be, became progressively more interdependent [2]. The domesti- maintaining keen hunting skills even when provided food and cation process likely began during the period when humans exhibiting a spectrum of behaviors ranging from untamable to ceased following wild herds of animals and adopted more highly affectionate pets [5,6]. agricultural lifestyles, particularly in the Fertile Crescent [3]. Cats spread to virtually all parts of the Old World, probably This change occurred 10,000–11,000 years ago and was made along trade routes between ancient civilizations. Despite their rapid spread, cats have remained amazingly similar to their wild ⁎ Corresponding author. Fax: +1 530 752 4278. felid ancestors (Felis silvestris subspp.) in form and function [2,6]. E-mail address: [email protected] (L.A. Lyons). The progenitor species of domestic cats remained compatible with 0888-7543/$ - see front matter © 2007 Elsevier Inc. All rights reserved. doi:10.1016/j.ygeno.2007.10.009 M.J. Lipinski et al. / Genomics 91 (2008) 12–21 13 Table 1 Genetic variation of cat breeds, random-bred populations, and wildcats Population N FIS HO HE Avg. allelic Avg. No. Alleles/locus richness alleles/locus range Abyssinian 40 0.13 0.45 0.52 2.44 4.29 2–7 American shorthair 13 0.09 0.57 0.63 2.90 4.59 2–7 Birman 28 0.10 0.43 0.48 2.31 3.82 1–6 British shorthair 28 0.11 0.56 0.63 2.96 5.82 2–9 Burmese 25 0.22 0.40 0.51 2.51 4.44 1–9 Chartreux 30 0.08 0.57 0.61 2.81 4.62 1–8 Egyptian mau 19 0.23 0.49 0.63 2.86 4.56 2–7 Exotic shorthair 40 0.12 0.55 0.63 2.96 5.91 2–10 Havana brown 11 0.13 0.41 0.46 2.28 3.03 1–6 Japanese bobtail 32 0.12 0.60 0.67 3.14 5.97 3–12 Korat 39 0.02 0.56 0.57 2.62 4.44 1–8 Maine coon 31 0.13 0.58 0.66 3.08 6.00 2–11 Norwegian forest 10 0.13 0.61 0.70 3.29 4.82 2–11 Persian 33 0.19 0.51 0.62 2.90 5.68 3–9 Russian blue 25 0.09 0.46 0.50 2.40 3.79 2–6 Siamese 32 0.10 0.47 0.52 2.47 4.06 1–8 Siberian 19 0.05 0.69 0.73 3.45 6.65 3–12 Singapura 24 0.12 0.34 0.38 1.98 2.82 1–5 Sokoke 14 0.07 0.41 0.44 2.14 2.82 1–6 Sphynx 27 0.10 0.58 0.64 3.00 5.59 1–9 Turkish Angora 14 0.15 0.56 0.66 3.08 5.21 2–10 Turkish Van 21 0.18 0.49 0.59 2.71 4.62 1–8 Breed average 25 0.12 0.51 0.58 2.74 4.71 Total/range 555 0.02–0.23 0.34–0.69 0.38–0.73 1.98–3.45 2.82–6.65 1–12 Brazil 26 0.08 0.66 0.71 3.40 6.82 2–13 Hawaii, USA 54 0.10 0.63 0.69 3.32 7.87 3–13 New York, USA 35 0.12 0.62 0.69 3.36 7.60 3–14 Texas, USA 30 0.10 0.66 0.72 3.50 7.18 3–12 Germany 45 0.14 0.62 0.70 3.41 8.39 2–15 Finland 32 0.08 0.61 0.65 3.22 7.00 2–13 Italy 47 0.1 0.67 0.72 3.56 7.8 3–14 Turkey 70 0.11 0.68 0.76 3.70 10.39 6–18 Israel 47 0.10 0.70 0.76 3.71 8.97 5–16 Egypt 27 0.11 0.68 0.74 3.67 8.26 4–17 Tunisia 17 0.04 0.68 0.69 3.38 6.45 2–12 Kenya 33 0.13 0.59 0.65 3.17 5.46 2–11 Sri Lanka 24 0.07 0.70 0.74 3.56 7.26 3–14 Singapore 29 0.10 0.65 0.71 3.47 7.68 3–14 Vietnam 20 0.08 0.63 0.66 3.27 6.42 2–12 Henan (China) 20 0.05 0.63 0.64 3.16 6.32 3–12 Korea 40 0.06 0.62 0.65 3.16 7.37 3–16 Random-bred avg. 35 0.09 0.65 0.70 3.41 7.48 Total/range 596 0.04–0.14 0.59–0.70 0.64–0.76 3.16–3.71 5.46–10.39 2–18 Felis silvestris caffra 10 0.30 0.57 0.76 3.98 6.55 2–10 F. s. tristami 5 0.11 0.60 0.49 2.41 2.60 1–4 F. s. silvestris 10 0.47 0.43 0.71 3.70 5.24 3–11 Wildcat average 8 0.29 0.53 0.65 3.36 4.80 Total/range 25 0.11–0.49 0.43–0.60 0.49–0.76 2.41–3.98 2.60–6.55 1–11 N, sample size; FIS, average inbreeding coefficient of an individual relative to its subpopulation; HO, observed heterozygosity; HE, expected heterozygosity; Avg. allelic richness, expected number of alleles in a sample of three diploid individuals, averaged over all loci. human agriculture. Gene flow between feral and tame modern lagged considerably. Although reasons to change the basic form cats, and between modern cats and their wild subspecies [7–10], and function of the cat were not as compelling as for other has not negatively impacted the role of cats as the principal small species, nonetheless, certain types of cats were artificially se- carnivore in human-dominated ecosystems. In fact, having a lected in various regions of the world. Interestingly, this “breed feral pool of modern cats surrounding the periphery of villages selection” was often based on aesthetics and involved simple and farms may have been advantageous for the control of pests traits of coat color, color patterns, etc. and less noticeably form or and associated zoonotic diseases. Therefore, the impetus to function. Cats did not lend themselves to become herders, change cats to suit certain human needs was much less than for workers of the hunt, or guardians, but their grace and beauty the other domesticated species and breed development for cats have always been obvious.