What is the Anatolian cat? This scientific review aims to define the Anatolian cat, its distribution area, genetics, and phenotype.
The review introduces to the natural cat populations and attempts to clarify the differences between natural cats and cat breeds.
Additionally, we will show how the natural selection and other evolutionary forces influenced and changed the cat populations all around the world.
Classification of Anatolian cats
Although taxonomists consider the domestic cats different from their ancestors and call them as separate species Felis catus (1), the genetic studies challenge this classification. Studies have shown that wildcats and phenotypic domestic cats are indistinguishable from each other at least in the Near East (2, 3, 4). This means that Anatolian cats are the Near Eastern wildcats and their taxonomic name should be Felis lybica lybica.
Anatolian cat is known by a number of misleading names: “sokak kedisi” (street cat), by breed names Ankara kedisi and Van kedisi (solid white Anatolian cats), (Turkish) Van cat, Cyprus Aphrodite Giants (Anatolian cats from Cyprus), Aegean cat (Anatolian cats from Greek islands) and Anatolian Shorthair (short-haired Anatolian cats).
The Anatolian cat (Anadolu kedisi) is a domestic cat, indigenous to Anatolia and nearby regions (4, 5, 6). Anatolian cats have a unique genetic profile and history, in relation to the region and its inhabitants.
Anatolian cats are present in Anatolia, Levant and nearby locations (Figure 1 and 2). The large portion of the Anatolian cat population is concentrated in urban areas.
The Anatolian cat is one of 8 natural cat populations
Undeniably the migrations, population interactions, and geographic isolation have left an imprint on the genetics of both humans and cats.
The spread of cats across the world is closely linked to human migrations (4). Anatolian farmers first brought cats to Cyprus (7) and Aegean islands, and not long after cats reached Europe (8). Cats were introduced to Asia and Africa continent likely by Middle Eastern and Levantine farmers (9).
Each cat ppulation in Europe, Asia, and Africa took its own distinct evolutionary path, and in thousands of years, these populations split from their Anatolian ancestors. These cats have established new populations in many geographic areas of the world. As a result, eight distinct natural cat populations evolved (5, 6). These cat populations are distinct, even though they are still regarded as the same species.
Natural cats are the populations of domestic cats that had evolved naturally, without human intervention. “Natural cats” is a more appropriate term than vague and weakly defined “random-bred”, “free-living”, “feral cats” or “community cats”.
We already know that all domestic cats trace their ancestry to wildcats that lived in Anatolia. But how is it possible that natural cat populations became so genetically different from Anatolian cats?
Separate populations of cats are produced through random evolutionary processes like the founder effect, explained below (Figure 3).
The illustration above explains how different cat populations are formed by “Founder effect”. Small colored circles in a larger circle display the genotypes of cats found in the original population – red and green are the most common and other color variants are rarer. A small number of cats migrate to other location (mostly red genotypes, just by chance), far from their parental population. These cats interbreed with each other, mutations occur, and so gradually these cats become a distinct population.
Natural cats should not be confused with cat breeds
It has long been believed that all cats are either breeds or a mix of several cat breeds. But the genetic research has revealed a very different story: natural cats are not breeds but distinct and ancient cat populations that predate all cat breeds (5, 6, 10).
[Natural] cats are the original populations from which the breeds developed, not a population of pedigreed cats gone feral
– Kurushima et al, 2013
All cat breeds were created by humans using natural cats in the last two centuries (11). Cat breeds do not occur spontaneously in nature, therefore such a thing like “natural cat breed” does not exist.
Natural cats from Europe and South Asia were founders of nearly all cat breeds existing today (12). For instance, Persian, Norwegian Forrest, Maine Coon and Siberian are descended from the natural cats of Europe (cats from Americas are of European origin; marked in green in a map, see figure 2). Siamese, Burmese and Russian blue have ancestors from South Asia (marked in color orange, figure 2). Some less popular breeds have different origins: Sokoke relates to West Indian cats, whereas the Turkish Van breed has ancestors from Anatolia (12).
Cats from Southeastern Anatolia Region. Photo credits: Çiğdem Köksal-Schmidt (Şanlıurfa), Mxfelix01 (Adana), Sobi (Aleppo), Hunbille (Mardin).
Genetic diversity of Anatolian cats
From cats to bacteria, every creature on Earth has been shaped by evolutionary forces. It took millions of years of natural selection to give rise to a lineage of small wildcats. One wildcat species changed its course of evolution when it chose to live alongside humankind 12,000-10,000 years ago.
The genetic legacy of these wildcats lives on in Anatolian cats. Cats from Anatolia and Levant have high levels of genetic diversity of all other cat populations which reflects the long history and population continuity in the region (6).
Genetic variation – the differences in genes between individuals. Each variation of a gene is called an allele. “Genetic variation” and “genetic diversity” are used interchangeably.
Lack of genetic differentiation between Anatolian cats from regions like Turkey, Cyprus, Lebanon, Israel and potentially Syria (not shown), indicate the outgoing gene flow due to migration among these populations (figure 4).
Non-Anatolian cat populations have reduced genetic diversity because of the founder effect (see figure 3).
Why genetic diversity matters? Genetic diversity is essential because it helps maintain the health of a population. Genetically diverse cat populations are capable of adapting rapidly to changing environments. Additionally, genetic diversity provides biological insurance against catastrophic consequences of disease epidemics.
Understanding the Natural and Artificial Selection
The source of genetic variation is a mutation, known as changes in the DNA code that alter existing genes or create new ones.
Many mutations are harmless – they do not have a noticeable effect on a cat’s phenotype or health. Mutations can be beneficial, for example, a gene that makes a cat resistant to a particular, while others are deleterious, like a gene causing kidney disease. Some mutations are beneficial and harmful: a gene may protect a cat from an infectious disease, but at the same time lead to cancer later on.
The natural selection is a process that acts upon genetic variation (mutations) present in a population. Natural selection is capable of reducing the frequency of many harmful mutations or eliminating them completely.
As a matter of fact, natural selection is responsible for healthier cats and “makes fewer mistakes” than human breeders, who just select cats for their appearance (13).
However, natural selection does not create perfect organisms. Recessive harmful mutations, such as diseases, disadvantageous morphological changes still occur in natural cats. Some of these “bad genes” can be passed down through generations, if a cat with such a mutation survives long enough to reproduce.
It is important to understand that every single trait found in cat breeds, be it a coat color, some morphological abnormality, and even an inherited disease – they all originated from the individual natural cats that became founders of cat breeds. To be exact, cat breeders did not invent something new: they just rearranged different gene combinations already present in a gene pool of cats used as starting material for their cat breeds.
The common argument is that breeding cats with unusual phenotypes, such as short legs, folded ears, without hair or with hair growth abnormalities etc. is justifiable because these are “natural mutations” (14). This is a typical example of naturalistic fallacy, the tendency to believe that what is “natural” is good (of course, breeders only use it as a moral justification for their choice to breed from disabled cats; on the other hand, cats altered by human selection are always regarded as superior to all other cats).
Just because a mutation occurs in nature does not mean that it is morally acceptable to breed animals with that mutation. From a cat welfare point of view, it is cruel to preserve the mutations that are a cause of disabilities in cats.
Anatolian cats may also carry deleterious mutations that sometimes that lead to visible phenotypic changes, such as shortened tail, dwarfism, and strabismus (crossed eyes) and similar (figure 5). While crossed-eyes are not attractive for cat breeders, cats with short tails and legs had been turned into “breeds”.
The phenotype of Anatolian cats
The morphology and phenotype of Anatolian cats are similar to that of other natural cat populations. Therefore, in general, categorizations of cats by superficial traits is neither objective nor scientific way to compare natural cat populations.
Anatolian cats exhibit sexual dimorphism: male cats are significantly larger, weigh more and are more muscular than females. They develop masculine heads and jowls, which make their heads appear round.
Traits like Brachycephaly (“flat face” in Persian breed) or dolichocephaly (“long head” in Siamese breeds), as well as a woolly coat with thick undercoat present in Persians and cats of European descent, are not seen in Anatolian cats.
Anatolian cats can be shorthaired or longhaired. They come in all usual color varieties: tabby patterns, including orange tabby, solid colors, like black and white; diluted: grey and cream; silver; calico and tortoiseshell in females, and all colors mentioned above in combination with white.
Clockwise from top left: mackerel tabby (wild-type), blotched tabby, mackerel with uneven stripes and spotted tabbies. Photo credits (clockwise from top left): Laurel Matt, Fatoş Böztepe, Ilyas Emir, Zülüf Ilk.
From the left: Mackerel orange tabby; blotched orange tabby and cream tabby (diluted orange). Photo credits (from left): Devrim Kılıç, Kevin and Sobi.
Solid black (melanism). The black fur may turn to reddish brown if a cat is deficient of tyrosine (15). Photo credits: Batu Aksoy (Izmir), Nükhet Barlas (Istanbul)
Many Anatolian cats display some degree of white patches in their coats (they are bicolored). Photo credits (clockwise from top left): Seyhan Ince, Talip Kilayıklı (Malatya). Sakazume (Istanbul), Ersin Akgöz.
Calicos and Tortoiseshells (females): black and orange; cream and grey. Photo credits (clockwise from top left): Zeynep Büyükçelen, Nükhet Barlas (Istanbul), Erkan Küçükşahinoğlu (Büyükada, Istanbul), Denis Senkov, Yunus Emre Dolgun.
Some coat colors common in other natural cat populations have not been observed in Anatolian cats (16, 17, 18). Colorpoint and its derivatives, chocolate, cinnamon, and diluted variants, seem to be limited to Far Eastern cat populations. These coat colors could only be acquired from hybridization with Siamese, Oriental type of cats or Persians.
Eye colors in Anatolian cats
Majority of Anatolian cats have green and amber colored eyes. Complete or partial heterochromia (two different-colored eyes) and blue eyes can be seen in white and bi-colored cats. Photo credits (clockwise from top left): Evren Özten (Istanbul), Kaya Güneş (Ankara), Fatoş Böztepe (Büyükada, Istanbul), Michel Berthaud (Istanbul), San Sebastian.
Behavior and personality
Anatolian cats express species typical-behavior observed in other domestic cats and small wildcats: they are carnivores, territorial and solitary predators who communicate with each other by rubbing, scratching and urine spraying (19, 20).
Anatolian cats readily exploit human environments. Due to supplementary feeding and access to food in waste bins, cats may form feeding groups or “colonies”. Living in high-density populations have downsides because this promotes disease transmission, inbreeding, fights, and even cannibalism (20, 21).
Not all Anatolian cats are tame. But if they are socialized from about the age of 2–7 weeks, cats usually tolerant and friendly to unfamiliar people (19). However, the personality and temperament vary significantly among individuals.
♦ Natural cats are neither cat breeds nor they are “mixes” of different cat breeds.
♦ According to the genetic analyses, there are 8 distinct worldwide natural cat populations, and one of these unique populations are Anatolian cats.
♦ Anatolian cats inhabit Anatolia, Levant, and surrounding areas.
♦ Anatolian cats look very similar to other natural cats from other geographical areas.
♦ Until recently, the Anatolian cat population has been subjected mainly to natural selection, and human influence was minimal. But with expanding cities, changing attitudes towards natural cats and policies that aim to make cities and everywhere else free of cats, the future of Anatolian cats is uncertain.
Cover photo: Başar Kocataş
Photos (with captions): Aysun İleriler (Anatolian cat) and Birgül Çildogan (Natural cats).
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12. Supplementary material: Table S3 (a) SNP assignment of cat breeds to random bred cat populations; (b) STR assignment of cat breeds to random bred cat population.
from Kurushima, J. D., et al. (2013). Variation of cats under domestication: genetic assignment of domestic cats to breeds and worldwide random‐bred populations. Animal Genetics, 44(3).
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