Anatolian Cat

Note: this publication is a manuscript. The complete and edited version of this research will be available in our website soon (in Turkish and English). 

What is the Anatolian cat?

This aim of this article is to introduce and define the ancient felines from Anatolia. Majority of ideas presented in this writing can be applied to all natural cats.

Anatolian cats/Anadolu Kedisi - ancient group of domestic cats living in Anatolian region and neighboring areas. These cats have unique DNA profile (1) and history, in relation to Anatolian region and its inhabitants.

Also called in these names: Ankara Kedisi, (Turkish) Van cat, Van kedisi, Sokak kedisi (Turkish: "street cat"), Aphrodite Giants, Cyprus cat, Aegean cat.

Natural cats– the populations of domestic cats (Felis Silvestris Catus (2)) that had evolved naturally, without human intervention. There are at least 8 distinct varieties of natural cats in the world (1, 3). Natural cats were exploited to create various cat breeds.

1. Origin & domestication

The Anatolian cat is a type of domestic cat, indigenous to Anatolia and nearby regions (1).
Figure 1: Feline family tree

Anatolian cats belong to the small wildcat species Felis Silvestris, which separated from other feline species 3.36 million years ago (4, Figure 1). All domestic cats, including those in Anatolia, are descended from at least five maternal lines of the Near Eastern wildcat, Felis Silvestris Lybica (5).

For a long time, it was believed that cats came from Egypt, and from there migrated to other parts of the world (6). It is not difficult understand why this belief persisted. Cats were a part of well documented religious cult in ancient Egypt. Ancient Egyptians left many artifacts portraying domestic cats and many more cat mummies. Since cats had a religious and cultural significance in ancient Egypt, there was no doubt that Egyptians were the first to domesticate them, or did not they?

The recent discoveries in archeology and genetics are already rewriting the history of the domestic cat.

Domestic cat did not come from Egypt or China

The study on thousand year-old Egyptian mummies revealed that mummified cats actually were very similar to the current populations of Egyptian street cats. It appears that a domestic cat came to Egypt later and before was tamed elsewhere (7).

According to one study, cats from unidentified species appeared in China earlier than in Egypt - about 5,300 years ago. These cats lived in early agricultural village and were fed by humans with grain based food (8). Media sources interpreted this study as an evidence, that cats were domesticated in China (9). However the species of cats found in ancient Chinese villages were later identified as leopard cats (10) - not F. S. Lybica species, from which domestic cats came from.

Humans did not domesticate cats; cats did it themselves

The oldest known finding, witnessing cat and human relationship, is ~9,500 year's old cat and human burial from Cyprus. (11) Early farmers from Anatolia brought the wildcat and other mammals to Cyprus by boat (12). It follows that cats and humans lived together in Anatolia for more than 10,000 years, since the beginning of farming (13). Wild cats at first were merely tolerated by humans. It was no more than a commensal relationship. It is very unlikely that Neolithic villages attempted to domesticate the wildcats roaming in their settlements. However in one particular situation, the wildcats could choose to “domesticate themselves”. It appears that preposition to tameness is a characteristic of many wildcat species, not limited to species which are the most related to the domestic cat (93).
Image the situation, when humans responded to a meowing of kitten, left by its mother, and began look after it. The cuteness and helplessness of the kitten triggered the nurturing feelings in humans. Cats, which had a contact with humans while kittens, behaved tame and were accepted as a part of Neolithic village life (50). 
The adaptation to living with humans was a very beneficial survival strategy for domestic cat ancestors. Tame cats had a stable supply of food: they fed on scraps from garbage left by humans and hunted rodents and other small animals, which lived in human settlements. Some cats were probably even fed by humans. This allowed the tame cat population to increase and largely replace the untamed wild cats in the areas they inhibited, although populations never became separate. 

Where are all these ancient Anatolian cats?
Illustration of 9500 year old cat and human burial from Cyprus  
Human and cat burial in Cyprus is an important discovery, which points towards the Anatolian origin of the domestic cat. Unfortunately, archeologists have not yet located any similar burials from Anatolia. Cat bone fragments are sometimes found in ancient settlements, such as Çatalhüyük (58) and Hallan Çemi (59), Aşıklı Höyük, Bademağacı and other ancient sites. But it cannot be known how these cats related to humans, wherever they were tame or wild. Inhabitants of Çatalhüyük welcomed the small predators and wildcats to their houses, because these animals helped to deal with mice invasion (61). Yet, the random bones of cats in Çatalhüyük (62) and other settlements tell us nothing about the place of cats in human societies.

It is possible that some wildcats were human companions, yet no evidence of their relationship survived. Cats could be buried in the area outside the settlements or some cats chose to leave human sites because they wanted “to find a quiet place to die” (55).

Feline symbolism frequently appears in Anatolian art from Neolithic period (60). Female figurines from Çatalhüyük and Hacilar are accompanied by felines, probably leopards and lions. It is possible, that some figurines may portray the domestic cat ancestors, as it seen in 7500 year old Hacılar example, where a woman holds an animal that looks similar to the cat.

While ancient art could be useful in a study of cat history, we should be careful not to take these portrayals at face value. The art featuring cats may be unrealistic and exaggerated. For example, the head of the cat from the later historical period, displayed in Ankara museum, has a very round face. It, of course, should not be used as a proof that “most of Anatolian cats had round heads” (56) - we know they do not (except for mature male cats with jowls).

Did the domestic cat came from Anatolia?

Archeology and history alone cannot answer the most important question: where the domestic cat came from? The genetic studies of recent cat populations already pointed out the species of wildcat that gave a rise for domestic cat lineages. And it is known already that this happened in the area vaguely defined as the “Near East”.

To reveal the exact location of cat domestication, we need to study the ancient DNA.

What is the ancient DNA? It is DNA extracted from ancient material, such as bones of felines which lived in ancient settlements and villages.

The team of scientists have been already working wıth mitochondrial DNA, extracted from remains of cats found in Neolithic settlement Bademağaçı (Antalya) and from other locations, such as Sagalassos, Aşıklı höyük, Demirci Höyük (Bronze Age). The primary analyses show that cats from these places belonged to F. S Lybica species (14). It may come as a surprise for many other researchers, who believed that F.S Lybica wildcats did not live in Anatolian region. Note, that F. S. Lybica wildcats are frequently confused with another wildcat, the Southern African wildcat (F. S Cafra). Southern African wildcat and F.S. Lybica are different species (5). Unfortunately many scientists still call the Southern African wildcats in F. S. Lybica’s name.

Compared to all other cats from various historical periods, the lineages of Anatolian cats were the most ancient, allowing us to speculate that the first steps of taming of cats were taken in Anatolia and probably Levant.

More recent lineages of domestic cats associated with Greek and Roman expansion, could come from the Middle East or Egypt. The domestic cat history appears to be correlated with historical movements of humans (57). The geography, relations between ancient populations and migrations have left an imprint in genetics of both Near Eastern humans and cats.

Are cats really domesticated?

Changes in behavior

Cats and humans coexisted for about 10,000 years. However during this period the domestic cats have changed very little from their wild ancestors (17, 13). The human influence on cat’s behavior and phenotype was minimal. Even in recent times many cats that live freely and avoid humans behave no different than wildcats. These domestic cats are called as “feral”.

Apparently, cats still are not domesticated animals (13). Scientists, studying the behavior of domestic cats, emphasize that the domestic cat behavior is very similar to that of wildcats (94, 95). Even those cats which grew up indoors are still able to revert to their wild stage if needed. Cats, whether kept indoors or living outside, do not lose their natural instincts. Jumping to high places, scratching, burying feces, chattering teeth at birds, hiding from strangers and feeling stressed in new places, being paranoid, killing small animals, fighting for territory etc. all these are the expressions of the natural cat behavior. By observing the behavior of wild cats we can learn more about the domestic cats and the other way around (95).

However, the domestic cat similarity to wildcats does not equal to them being aggressive, fearing of humans and behaving in other inappropriate ways. Domestic cat ancestors were more successful than other feline species because of their ability to adapt to their environment. Tameness and friendliness towards humans do not come naturally – this is a learned behavior. A cat in order to become a “good pet”, needs to be socialized (96). Socialization of freely living cats in urban areas is not difficult. Kittens, which did not experience a harm and abuse that could make them distrust humans, are given food and had an opportunity to come into close contact with humans, will likely grow up to friendly and affectionate adults.

Natural cats are ruthless predators. They can survive on their own without help from humans.
Photo by Zülüf Ilk

While the natural cats are still are very similar to its wild ancestors, there is one group of domestic cats which are more or less domesticated. These cats are man-made breeds, such as Siamese and especially Persian (16). In the last century, humans began to manipulate the cat’s appearance and behavior. Breeders were not interested to preserve the traits that made a cat such an excellent survivor; they aimed for exaggerated body and fur types, rare phenotypic mutations (for example: folded ears) and docility.

Changes in phenotype 
The wildcat from which the domestic cat originated, was a shorthaired tabby. The mackerel tabby color (see the figure) is the most ancient and a primary color of the cat. (18, 42). So how other colors arose? From random mutations in wildcats. However these variants were not favored by the natural selection: they were very rare and offered no evolutionary advantage.

The situation changed when cats started to live near to humans. The rare color and longhair mutations were noticed by humans. Cats with unique looks stood out from the rest of shorthaired tabbies. Humans likely preferred these different looking cats, and indirectly selected them (89). Of course, the selection was unintentional: humans did not breed cats purposely (19). Ancient communities were probably more tolerant to the presence of different looking cats, and some probably had superstitions and magic beliefs about these cats. Whatever the reason was, unusually looking cats were encouraged to stay in the villages.

Humans were indirectly responsible for continuity of majority of phenotypic mutations in cats (89). The mutations influenced by humans are present in nearly all modern populations of cats: melanism (black), dilution (grey, cream), long fur (from SH-2 haplotype), white spotting, classic/blotched tabby, orange/red tabby, and dominant white (20).

Anatolian Cats = wildcats?

The line between domestic cat and a wildcat is blurred, because domestic/tame cats were never isolated from its wild siblings, F. S. Lybica. The ongoing interbreeding between the wild and tame cats, prevented the separation of the wild and domestic lineages (24). This would explain why domestic cats and Lybica-type wildcats from the Near East, are indistinguishable from each other genetically (5, 25; see Figure 2). In short, F. S. Lybica and Anatolian cat could be the same cat. This could change the understanding both of the F. S. Lybica wildcats and the Anatolian cats. This also would force to rethink the methods for their conservation. Currently F. S. Lybica species are not considered to be endangered. (51)

Figure 2. The wildcats (F. S. Lybica) and domestic cats from the Near East (Anatolia, Levant, and Mesopotamia) are identical genetically, shown in brown color. However the domestic cats from Europe, Asia and Mongolia are distinct (Driscoll et. al, 2007). (5)

2. Natural cats – disliked, pitied and misunderstood

Domestic cats are very successful species. They, with a help from humans, populated nearly every part of the world. Domestic cats are able to survive in many different environments. Unfortunately, the success of domestic cats, became a threat to other species of wildcats: European wildcat (21), Scottish wildcat (22) and South African wildcat (23). Domestic cats can interbreed with different species of wildcats and produce the fertile hybrids. This makes conservationists and biologists to worry.

In most of Western world, the natural cat populations are not regarded in a positive light. They are blamed for killing protected species, spreading diseases and judged as unwanted, homeless animals – “a burden on society”. In contrast, man-made cat breeds are seen as valuable and desired. Nearly all the scientific literature emphasize the negative aspects of natural cats (63). The research about natural cats is nearly always about the population control, toxoplasma gondii and how freely living cats damage the wildlife. No effort is given to understand and define these diverse cat populations, or improve the aspects of welfare which go beyond the sterilization and shelter-adoption system.

Natural cats are controlled by sterilizing and by locking them into shelters. Every year millions of cats are killed in USA shelters (64, 65). In Australia and New Zealand, the domestic cat is seen as a pests and aggressively exterminated (66, 67). In some Asian countries such as South Korea (68) and in Africa (Madagascar (69)), natural cats are both kept for companionship and killed for meat. No place in a world is a “safe heaven” for a domestic cat.

The attitudes towards natural cats are different in Turkey: locals do not see cats as ruthless predators, euthanasia is unacceptable (70) and there is no concern about how cats may affect the wildlife. A cat is regarded as a victim of car accidents, diseases and human activities. Population control is encouraged in order to “reduce suffering” because freely living cats thought to be “abandoned and left on the streets by humans” (74) and unable to survive on their own, because they are “domesticated”.

Cats co-existed with humans for thousands of years in Anatolia and most of other Eastern Mediterranean countries. They were and still are a part of local cultures.
Cats in Anatolia traditionally were not kept as pets in a traditional sense. It was rather a commensal relationship or “semi-ownership”. Many cats developed the close connection with Anatolian locals by whom they were fed and looked after. The good examples of this type of human and cat relationship was portrayed in a documentary movie about cats of Istanbul called “Kedi” by Ceyda Torun (76).

The current welfare model practiced by animal activists in Turkey, was inspired by Western countries mentality and beliefs. The internet played a huge role in changing perceptions about cats in Turkey. One of the first sites about cats, actively promoted neutering campaigns, shelters and even encouraged segregation of cats based on breed labels.

The globalization and the popularity of Western welfare model among welfare organizations and policy makers, could mean that Anatolian cats will be further devalued, in result their lives will likely get only harder while their numbers will continue to decline.

“Sokak Kedisi” stereotype

The Anatolian cats are commonly called as "sokak kedisi", or a street cat in Turkey. "Sokak kedisi" describes a homeless or a stray cat which lives on the street. This term carries a negative image, while a term “breed” is associated with a higher value and prestige.

"Tekirim diye beni kimse istemiyor" - "Nobody wants me because I am a tabby"
A poster from one adoption ad. 

Anatolian cats are mistakenly regarded as “mixed” with cat breeds, although a few are said to be “purebreds” (26). How these breeds are identified? Well, it is done mainly by judging a cat by its color of fur (53) For example, majority of white cats are said to be Van Kedisi (30) or Ankara Kedisi (29), black cats – Bombay breed (28), grey cats – Russian blue, and colored longhaired Anatolian cats – Norwegian Forrest’s (52), Siberians or Maine Coons! Tabbies, for some reason, are universally thought to be a “mixed” (melez/kırma) breed (26). It is not clear where the tendency to classify the natural cats into breeds based on the color of their fur, came from. These first websites in Turkey about cats, and later their replicas, popularized the cat breeds and were responsible for misinformation too (the belief, that cats with colored cats, especially with tabby patterns are “mixed breed”). Breeder’s publications were and still are the main sources of information about cats. Apparently, cat breeders and registries shaped people’s perception about natural cats in Turkey and everywhere in the World.

Although a white and a tabby cat may look very different from each other, the look is misleading. Often, a white and a tabby cat will be siblings or close relatives. The difference between a white cat and a tabby cat is that white cat has a white allele which covers up its color. Majority of white cats are heterozygous, it means they have both white and another color in their genes, just because white allele is dominant, the other color is hidden (32). So the white cat can be a tabby even if it appears completely white.

Grey cat looks grey, because of mutation in MLPH gene (31) which lightens the black color. These single color genes do not make cats to the different breeds.

We do not say that humans with black hair and those with blond hair are different “races” or that one with blond hair is purer race than another with black hair. This would be dismissed as a racism. But why it is seen completely normal to judge cats by the color of their fur? Even scientists are guilty for racist generalizations about the cats. For example, one study claimed that calico, tortoiseshell, black and white cats were more aggressive based on anecdotal reports from their owners (33). There is no correlation between the color of fur and behavior. One notable exception is when a cat carries a white allele which also makes a cat deaf, but again the altered behavior is caused by secondary factor (deafness) not because it is white (not every white cat will be deaf).

What about the breeds, which said to be occurring “naturally” in isolated populations? (34) Cat breeds do not just happen out of nowhere on the streets. Breeds in order to exist, must be invented by cat breeders. Breeders exploit the rare phenotypic mutations found in natural cats. No matter what cat breeders and their registries are saying, there are no “natural breeds”. A cat is not a breed, if it did not go through the strict artificial selection.

Keep in mind, that breeders do not create something new: they just rearrange different gene combinations together. All these gene combinations (variations), whether rare phenotypic mutations, colors or diseases come from one source: the natural cats. Natural cats are the founders of the breeds (3, 35). Even the most bizarre looking breeds, like hairless cats, are not some kind of laboratory experiment, but a rare gene mutation, accidently discovered from a street cat, and made to the breed by an ambitious breeder.

People believe that cat and dog breed histories are similar, so they assume if stray dogs are mixed breeds, so the same must apply for cats too. Does the claim that Anatolian cats are mixed with breeds make any sense?

Think about this. Lots of cats from controlled breeding programs and with recorded origins (otherwise how can we be sure it is a breed?) must be left on the streets. Thousands of them. But we know that cat breeds are just a tiny minority, according to some estimates not more than a few percentage of all cat populations in the world. Also it is highly unlikely that lots of people would leave their expensive pets on the streets. Even if many breed cats get abandoned, these cats must be intact and be able to compete with natural cats. Only if breed cats somehow manage to outnumber and replace Anatolian cats, only then Anatolian cats would become mixed with breeds. Of course, this situation sounds like a conspiracy nonsense. Unfortunately, this type flawed thinking appears to dominate the public opinion in Turkey, because some people believe that cat breeds are widespread on the Turkish streets. This wrong belief is also strengthened by anecdotal claims and guesses that some cats were pets and later were abandoned by humans - none of which can be verified.

Geneticists, who studied natural cat populations, disproved the idea that natural cats originated from the breeds. In contrary, the natural cats around the world are "the original populations from which the breeds developed” (1, 3, 35, 36). So the ancestors of cat breeds were “street cats”, but the “street cats” did not originate from the breeds. Some cat breeds are less than century ago, as for example Persian and Siamese, however majority other breeds were created in 1950’s using Persians, Siamese and European street cats (8). The natural cats however existed long before any cat breed!

There is even a DNA test (37) developed for those who strongly believe that a cat they adopted from the street, is some kind of exotic breed, and want a confirmation for their belief. But such a test will probably be a waste of money, because natural cats are not breeds, so they cannot match the genetics of small and limited populations of breeder’s cats.

In conclusion, the Anatolian cats did not come from abandoned or lost pets. They are not a human creation. They are descendants of ancient cat lineages, which lived in Anatolia for many thousands of years.

Natural cats - what are they?

Until recently, natural cats were thought to be “mixed” and uninteresting from the evolutionary point of view. It seemed like, no matter where they come from, all these cats were almost the same - just mixed “street cats”.

Scientific community had a little interest to study the natural cats, unless it was about population control, parasites or the damage cats cause to the wildlife. Nevertheless some interesting discoveries were made when geneticist began to research the cat breed origins. They found out that the domestic cat had a far more complicated evolutionary history.

Much to our surprise, the natural cats from different places of the world were actually different. (3) And these cats had nothing to do with breeds, other than cat breeds could trace back their origins to some natural cat populations.

Natural cats correspond to separate groups, in relation to their geography (see: Figure 3). The geographical and genetic differences reveal that natural cats do not really breed randomly as believed. Hence, we do not use the term “random-bred”. Natural cats usually breed with other natural cats living nearby. Their gene flow is restricted by geographical barriers and the distance. The larger the distance, the more different cat populations turn out to be.

Although the domestic cats are different at the genetic level, they still have a very similar appearance. This has to do with the fact that their most recent common ancestor lived less than 10,000 years ago. Besides there was no selective pressure on cat’s phenotype. It does not mean that these cat populations did not acquired adaptations over time, just the changes had minimal influence on their phenotype. As we can see from the map (Figure 3), the distribution of particular grouping of cats have nothing to do with political borders, and cat varieties often overlap.

There are about 8 domestic cat varieties: Anatolian & Levantine, Middle Eastern, Western European, Egyptian, Indian Ocean region, Indian, South Asian and East Asia (4). Western European cats can further be divided into 4 more separate groupings (not shown).
Figure 3. Natural cat populations of the world. A map was made according to data of microsatellites (inverse distance function) by L.A Lyons and J. Kurushima (4)). This map shows the populations of natural cats and their distribution worldwide. Middle East does not mean "Persian cat", South Asia is not about "Siamese" and Egypt is not about "Egyptian Mau", and Anatolia includes all native cats with long/short fur in variety of colors.

The distribution of varieties natural cats had a lot of to do history and human migration. Anatolia and Eastern Mediterranean regions were related historically and culturally, so cats there have similar genetics. Centuries ago, European immigrants brought cats from Europe to Americas, Australia and New Zealand –this is why cats from Western World are still “European”. Far East was largely isolated from the West, and this shows up in their cats too.

3. Phenotype of Anatolian Cats

Are longhaired and shorthaired cats different?

White cats are perceived as Ankara and Van breeds in Turkey. However some individuals have recently begun to seek recognition for the third breed, as they call it the “Turkish shorthair” (Kısa tüylü Türk kedisi). They even measured the shorthaired Anatolian cats to create their “standard” (38). Nonetheless this idea is not very original, because one cat breeder’s registry had already promoted the shorthaired cats as a breed under the name Anatoli (39).

Let’s make it clear: it does not matter if a cat is longhair or shorthair - Anatolian cat is NOT a breed.

Obviously, a cat with a long fur has a longhair gene. Let’s call it in a letter “L”. This gene “L” is weaker (recessive), because a cat must inherit two copies of it in order to be longhair (see Figure 4). This is why longhair cats are uncommon compared to shorthaired ones. Fortunately, the longhair gene is not likely to die out, because many shorthair cats carry one copy of that gene. It is possible to get longhaired kittens from two shorthaired cats, if each mother and father have a copy of “L” gene (see Figure 5). Because many Anatolian cats have a hidden “L” gene, many litters will consist of both longhair and shorthair kittens.

Figure 4. Mendelian inheritance of long fur. The short-haired cats often carry a long-haired gene and when mated with longhaired cat or another carrier, may have long-haired kittens.

Figure 5. Mating between shorthaired and longhaired cats do not produce intermediate (something between short and long fur) variants, as believed. The kittens born to one or two short-hair parents may have the fur similar to those cats having two long-haired parents.

Longhaired Anatolian cats

The term “Angora” or “Angora phenotype” is used in scientific terminology to describe a mutation of the long hair in animals (40, 41). We also use the alternative name “Angora” for Anatolian cats with a long fur.

All domestic cats came from a shorthaired and tabby wildcat. Other colors and a long fur mutations appeared when wildcats became tame and started to live near to humans. These mutations persisted through generations of cats, because of indirect human influence.

The gene that makes cats’ longhair, has been identified (44). FGF5 (Fibroblast Growth Factor 5) gene regulates the growth of hair in humans (41) and fur length in dogs (45, 46). A mutation in FGF5 gene resulted to a long fur in cats (44, 47). This ancient mutation is known as c.475A>C is found in all cats around the world (48). It was discovered that this mutation came from one rare short-hair variant, called SH-2 haplotype (48)

"Although long-hair is thought to be a simple trait, the actual hair length of a long-haired cat is highly variable"(48); So the length of hair and their thickness are slightly different in each cat. Several factors determine the appearance of the fur: individual cat's genes, cat’s health, age and nutrition. The fur of longhaired cats tends to be significantly shorter in summer season. Many cats will shed a lot of fur and will look similar to shorthaired cats.

The cat fancy uses the term "semi-longhaired“to any cats which do not have a fur comparable to Persian breed (34). This is wrong, because the Persian fur is an abnormality. Why other cats are measured up to Persians? The fur of Persian is an exception. The correct term to describe the Persians would be the breed with “abnormally long” fur, as suggested by UFAW (43).

To recognize the longhaired cat is not difficult, since its fur is obviously longer, and this is most apparent by looking at the cat’s tail. Such a cat carries two copies of the longhair genes. So this cat should be called as longhaired, regardless the length of its fur.

Identifying the Anatolian cat

Anatolian cat does not have one type of look or one type of personality. The Anatolian cats look similar to other natural cat populations. 

The Anatolian Cat comes from Anatolia, Levant and nearby areas.
Cats living on the streets of Turkish cities are Anatolian cats.
Anatolian Cats do not display exaggerated traits, present in breeds 
* Anatolian Cats with a long fur can be alternatively called as “Angora cats”, but they are not different from the shorthaired cats. 
The size and body weight is within range of domestic cat. Male cats are bigger, weight more and are more muscular than females (49).They develop masculine heads and jowls, which make their heads appear round.
* Anatolian cats come in all usual color varieties: tabby patterns, including red tabby, solid colors like black and white; diluted: grey and cream; calico and tortoiseshell in females, and all colors mentioned above in combination with white. 

Eye colors of Anatolian Cats: green, amber and its shades, odd-eyed (partial and complete heterochromia) and blue eyes in white and bicolored cats.
Photos by Evren Özten, Kaya Güneş, dimashoo, Michel Berthaud and Fatoş Böztepe

Clockwise from top left: wild type tabby (mackerel), blotched tabby, mackerel tabby with uneven stripes and spotted tabbies. 
Photos by Laurel Matt, Fatoş Böztepe, Ilyas Emir, Zülüf Ilk
From the left: Mackerel orange tabby; blotched orange tabby and cream tabby (diluted orange)
Photos by Devrim Kılıç, Kevin and Sobi

Grey tabbies
Photos by Enrico Löffler, penjelajah.jepang, Nükhet Barlas
Silver tabbies
Photos by fati-foto, Fraggler Ocking Troodos, Levent Deniz
Solid white and grey
Tortoiseshell (females): black and orange; cream and grey.
Photos by Emre Zaimoğlu, Denis Senkov, Yunusemre Dolgun, Erkan Küçükşahinoğlu
Calico (females): Tortoiseshell with white
Photos by Zeynep Büyükçelen, Nükhet Barlas, Belkıs Eksioglu, Burak Sezer
Solid black (melanism). The black fur may turn to reddish brown if a cat is deficient of tyrosine.
Photos by Batu Aksoy, Nükhet Barlas

Bicolored (combinations with white)
Photos by Kazuko Sakazume, Talip Kilayıklı

Characteristics not observed in Anatolian cats:

Ø Foreign (slender), oriental body type in mature cats. Malnourished, sick cats tend to be appear very slim however it is not a normal condition. 
Ø Excessively round and short body type with brachycephalic head indicates Persian out-cross. 
Ø Very large, close set ears and a long head (dolichocephaly) - the traits of the Turkish Angora breed. 
Ø Pointed colors, chocolate, cinnamon and its diluted variants, which come from mixing with Siamese and Oriental type of cats or Persians. Pointed colors and its derivatives originated in Asian cat population, and are not found in Anatolian cats (99, 100, 101). 
Ø Dark blue eyes in white cats in most cases show that a cat hides pointed color under its white fur (32). 
Ø Woolly coat with thick undercoat comes from outcrosses with Persians.

Behavior and Personality of Anatolian Cats

When people think about the cat breeds, it is not only about the different look. They believe that cat breeds have generic and predictable personalities.

The descriptions of certain breed’s characters are that much accurate as astrology signs, explaining the complex personalities of human beings. In other words, they are vague generalizations, guesses and often will match to almost any cat.

Only certain cat breeds have traits which are more or less predictable: like Siamese, being more outgoing and demanding, Persian – a slow and inactive cat and a Ragdoll, selectively bred for docility (103).

The character traits of Anatolian cats are variable and it would be unfair to attempt to generalize them. Every cat has a unique personality shaped by its genes and environmental influences.

The cat behavior is not correlated with its fur color. There is no logical explanation of why a cat’s color would have any influence on cat’s personality and character. However in cases of disability may influence cat's behavior, as for example with deafness.

Natural cats can be as much as affectionate as a cat from controlled breeding. If only we assume that it is bred for its temper, because not all pedigreed cats are well-behaved!

The socialization has a strong effect on feline friendliness. In the areas where cats receive little attention, rarely fed by humans, these cats are likely to display more wild behavior, distrust and anxiety towards humans.

This is why the friendliest cats are found in larger cities where its inhabitants have a tendency to care for their cats; food and treats act as a powerful reward, drives and encourages the affectionate behavior. The friendliest cats get the most attention and benefit from human care, in result they are more likely to pass their genes, at the same time a preposition of tameness for their future kittens. This is an example of indirect selection by humans, unconsciously practiced since the beginning of human civilization. Currently the situation has changed drastically, because selection now moves in the negative direction. Where neutering the natural cats became a norm, the friendliest cats became an easy target. In contrast, untamed cats, which are not good candidates for sterilization, because they are difficult to catch in a first place, successfully reproduce. Cats, which instinctively fear and avoid humans, will have kittens behaving similarly, making the whole population of community cats less human-friendly.

There is no reliable way to predict personality of a cat. Only the friendliness or tameness are exceptions. Certain regions in cat genome (77), particularly those associated with "stimulus-reward learning" are probably responsible for tameness in cats. "Tameness or domestication" genes are probably natural variations present in natural populations, not spontaneous mutations (78). However it appears that contribution of "domestication genes" to cat behavior is quite modest. The domestic cat is still closely related to its ancestor, a wildcat. Thus by allowing the friendliest cats to breed, the chance that the kittens will have a similar behavior, is likely (16).

There are many factors that will determine the friendliness of the cat:

1. Age of separation from mother: In one study (79) kittens were separated from their mothers at 2 weeks old and others- at 2 and 3 months, and then their behavior was compared. Kittens taken from their mothers younger than 2 months old, had significant behavior problems when grown-up: fearfulness and aggression directed towards humans, anxiety, impaired learning ability. Some of these cats even developed asthma-like syndrome (79). 2. Socialization (or lack of it). The amount of handling in first 8-12 weeks had a long lasting effect on cat’s friendliness (80,81). However a minimal interaction between humans and kittens was ineffective. Some cats were aloof even when properly socialized (82). The genetic factors may be stronger than socialization in some cases. Cats successfully will form social attachments to other animals, like dogs. It is important that a cat would get introduced to a dog early (83). Cats which encountered a dog for the first time not older than 6 months, had the most harmonious relationships, the least aggression and were able understand dog’s body language well. The same could be said about the dogs which started to live with the cats at the age of 1 year or less (83). The ability to silence predatory instinct in cats had been tested with rats. Kittens which grew up with rats, considered them as their siblings rather than a meal (84). 3. Mother: kittens imitate behavior of their mother. For example, they copy her predatory behavior and their choices of prey will depend on their mothers preferences (84,85); kittens prefer the food their mother selects to eat, even when it is a less satisfying choice (mashed potato instead of meat pellets) (86). 4. Father: kittens sired by friendly father were also more affectionate and less-stressed. They did not fear strangers (81,87,88). The father's behavioral traits could be inherited. There reason for this could be the fact, that mammals inherit more genes from their fathers than mothers.

How objectively can you judge the cat’s behavior?

Humans tend to anthropomorphize animals (71). In other words, humans attribute human characteristics and emotions to cats, and forget that the cat sees the world very differently. These are two examples, how humans project their own feelings to their cats: First example, cat suddenly begins to urinate outside its little box. The owner thinks, that a cat does it out of anger or wants to revenge for something. But the owner is wrong: the urination outside litter box is caused by the medical condition (73). Second example, while playing, a cat scratches owner’s hand. The owner thinks his cat is aggressive. However it is an owner’s fault: Play-related aggression is common in cats which were allowed to play with owners hands (97)

People often over-generalize the large group of cats based on what they observed in their pets behavior. "My cat is very fond of me, so all (…) cats are like this", "they are like dogs" – all these are nothing more than personal opinions. Of course, these observations may be true about some cats, but certainly will not describe all Anatolian cats. In fact, we will be quick to learn that if we take another cat to our house, it will be nothing like the previous one we have had before. It is important to understand that the natural cats have very distinct personalities and all are individuals, not to be judged by one superficial standard.

4. The future of Anatolian cats

Figure 5. Examples of phenotypic mutations in Anatolian cats: bobtail, short legs and cross-eyes. 

Natural & Unnatural selection

Every natural cat population is shaped by evolution. The Anatolian cats are ancient, so they accumulated a lot of variation, in other words - unique mutations. 
Many mutations are neutral – they do not have a noticeable effect on cat’s phenotype and health. Some mutations are beneficial, for example a gene that makes a cat resistance to particular disease. Some are harmful, like a gene causing kidney disease. Some genes can be both good and bad: a gene may protect a cat from an infectious disease, but the same gene may also cause it a cancer later.

The natural selection, a beautiful metaphor for a really cruel natural process, serves its purpose in eliminating unfit individuals with harmful mutations, "assuring" the continuity of the population.

Unfortunately the natural selection is not perfect. Detrimental mutations, such as diseases, disadvantageous morphological changes still occur in natural cats. These “bad” genes be passed down through the generations, if a cat with such a mutation survives long enough to reproduce.

There are rare examples of Anatolian cats which are born with unusual phenotypic mutations, such as shortened tail (bobtail), short legs and crossed eyes (Figure 5). Note, that short tails and legs are disabilities and disadvantages for natural cats. However the cat breeders in West exploited these detrimental mutations to create various cat breeds. From a cat welfare point of view, it is cruel and unethical to perpetuate disabilities of cats and promote them under breed names. 

How unnatural selection could harm the Anatolian cats? 

Natural selection is responsible for healthier cats and does significantly less mistakes than human breeders, who just select cats for their appearance. The natural selection actually reduces the frequency or even eliminates the harmful mutations. But the Natural selection can work only if the population of cats is sufficiently large (90).

Human driven evolution, for example by neutering freely living cats, is called unnatural selection (91, 92) . The unnatural selection will possibly have negative impact on future generations of Anatolian cats. If gene pool of Anatolian cats becomes small, the Anatolian cats may then suffer from various diseases which otherwise would occur in low frequency when the population was large. 
Cats, which are able to escape the neutering campaigns will continue to breed, however if these cats turn out to be carriers of rare and debilitating diseases, the whole remaining population of Anatolian cats will eventually become unhealthy (98). A population plagued by various diseases and lacking variation that is essential in order to adapt to its environment, is destined to extinction.

Welfare: many problems and wrong solutions

Photo by Birawar najm (Iraq)
The future of Anatolian cats is uncertain. It largely depends on the public understanding and governmental policies.
The current welfare policies of Anatolian cats are mainly focused on population control, sheltering, rehoming and feeding. Many cat lovers think about a cat welfare as a simple matter, that all we need to know about the welfare is already researched and solutions are easy.
Cat welfare decisions are often reduced to human emotions and feelings, which are also projected as the needs of the cat (71, 72). For example, according to human thinking, being homeless is a bad thing, so they attribute this feeling to the cat, imagining that a cat also has a similar desire to have a home (75). This leads to various consequences: harmless, such as making houses for the cats living on the street, and harmful, such as collecting cats to the shelters or hoarding them at home. Currently most of decisions about cat welfare do not involve science. However, a welfare which is not based on science, could actually do more harm than good for cats. Cats do not think like humans do and their problems and needs cannot be explained by “what humans feel is best for them”.

The concerns of Anatolian cats 


Ø Herpes infections 
Over 90 % of cats around the world are infected by Feline Herpesvirus Type 1(FHV-1) (102). The Anatolian cats are not an exception; they are likely to have a high rate of FHV-1 infection. Once a cat becomes infected with herpes infection, it may shed virus throughout its lifetime. Infection reoccurs, if a cat has a lower immunity, and stress is the most common cause. Cats can be additionally infected with secondary pathogens including calicivirus, Chlamydophila felis, mycoplasma sp., and/or bacterial pathogens (103). These conditions are treated with antibacterial drugs. Herpes infections however, do not respond to antibiotics.

Kittens infected by FHV-1: mild and severe cases.
Photo by Asli Kodan

FHV-1 infection is most severe in young cats and kittens. Herpes infection usually appears in kittens, when maternal antibodies decline (8 weeks of age). Kittens exposed with herpesvirus first time will have conjunctivitis (eye infection) with upper respiratory tract disease (104). If infection is left untreated, it may lead to blindness. Thousands of Anatolian kittens will become blind and suffer if infection will be left untreated. This tragedy can be prevented with antiviral drugs. The most effective antivirals are famciclovir and cidofovir. But they are effective only if applied often, a few times a day for a longer period of time that is difficult and impractical for cats living outside. These drugs are expensive. We need support the research that would help us to find better and more effective treatments for herpes infections.

Ø Deafness

Deafness may happen in any white domestic cat, but why it is an issue of Anatolian cats? Because people purposely breed the white cats thinking these cats are different from the other “street cats”. White cats are said to be either Van Kedisi or Ankara Kedisi “breeds”. The breeding of white cats is institutionalized and supported by Turkish government. 

Inherited deafness is very rare in colored cats, but many solid white cats tend to be deaf (33). White cats occur naturally: these cats have a lower chance of deafness, because they tend to be heterozygous, and deaf cats most often will not survive outside.
The dominant white gene makes a cat white, but also interferes with cats hearing: all homozygous white cats are deaf, while those carrying a color allele have a significantly lower rate of deafness (34). Breeding deaf cats or any white cats with unknown hearing status, is strongly discouraged. Breeding hearing (as confirmed by electrodiagnostic test) white cats with the colored ones, reduces risk somewhat, but does not prevent the deafness in kittens completely. 
In general breeding the solid white cats is risky and unnecessary. Alternatively, people should prefer cats with some color. Those cats with a white spotting gene look mostly white, but they are not deaf. 
The official breeding programs in Van kedisi evi and Ankara (Keçiören) refuse to acknowledge that deafness problem exists at all, by citing anecdotal and unverified claim, that only 2-3 % of their cats are deaf. It should be discouraged to breed white cats, not only because of high risk of deafness, but also because it is unscientific and a very racist idea. 
White cats not only suffer from deafness but they are also susceptible to skin cancer. White cats should not stay under the sun for long periods of time. The sunburn may cause squamous cell carcinoma, particularly on its head, in areas where fur is sparse. White cats are about 13.4 times at risk compared to colored cats (32). 
Remember, deafness is a disability. Deaf cats should be kept indoors or in protected enclosures only.

Ø Obesity: Obesity is very serious problem. In humans, it associated with many diseases, such as diabetes, heart disease and cancer. Neutered cats of both sexes tend to be heavier than cats intact (7). Energy requirements decrease after spaying or neutering, this is why owners of these cats should monitor food intake and calories to prevent obesity and with it related complications (35a). Freely living neutered Anatolian cats suffer from obesity too. It is very difficult or maybe impossible to control how much and what these cats eat. 

Ø Inherited diseases – the developments in feline genetics and DNA tests were not intended to benefit the natural cats. The genetic research was done on breed cats and to solve their health issues. While natural cats may use the same genetic tests, just like breed cats, because they may have similar diseases, the benefit is limited. The diseases present in breed cats are usually rare and could be already eliminated from the natural cat populations. Besides, most of owners of natural cats will not breed their cats, therefore the detection if a cat carries a bad mutation, has little value for them. Freely living cats, of course, gain nothing from the genetic screening at all. So the genetic tests exist benefit the cat breeders.

The natural cat populations are regarded as healthier than breeds. However the natural cats are constantly evolving, not necessary in a good way. Natural cats, including Anatolian cats, are negatively affected by unnatural selection, therefore the frequencies of disadvantageous genes could increase and the diseases may be different from those which affect breeds. While the genetic tests are already helping to get rid of diseases from breed cats, the genetic health of natural cats is neglected. The natural selection may not help for natural cats either, because when its population gets smaller, the natural selection is unable to function properly.

The powerful gene editing technology CRISPR currently is not used in feline research. However it is a matter of time when laboratories will start to offer this service for breeders. In the near future, breeders may artificially alter the genes of their cats. Breeders could eliminate various diseases and even produce superior individuals that selective breeding alone cannot achieve. Only if people become interested to support the genetic research for natural cats, for example, via crowdsourcing, only then we can expect that the achievements in genetics will not be used by privileged few, but for a benefit of all cats.

Blood type incompatibilities in Anatolian Cats

The risk for blood group incompatibility is very high in Anatolian cats. If a cat with a blood type B receives a blood from cat with a type A, this can cause haemolytic anaemia and even death. Blood typing of donor and recipient cats must be done before blood transfusion. If females having the blood type B mate with males with type A or AB, their kittens may die due to condition known as Neonatal isoerythrolysis. Many Anatolian kittens probably have died this way, however the reports on this condition are very rare, maybe because the veterinarians and general public have a limited knowledge about feline blood incompatibility. In controlled breeding, blood typing must be performed before mating – Female with B type of blood should not be bred to males with A and AB blood types. Kittens born from cats with A-B mismatch, should be removed from their mother at birth and fed with milk replacement for 24 hours. However it does not guarantee they will survive. In freely living Anatolian cats, to prevent kitten deaths due to A-B mismatch, is be very difficult.


* Support veterinary medicine and genetic research on natural cats by spreading awareness and participating in crowdfunding. We need better and more effective treatments of diseases that affect the natural cat populations. FHV-1 is one of these infections which ruin lives of thousands of hundreds of kittens by causing blindness but lacks effective treatments.

* Discourage the people and institutions from breeding white and deaf cats. Cat's ability to hear is more important than uniformly white fur.
Demand and support development of alternatives for surgical neutering. We need less invasive, less time-consuming, and less damaging options to control the reproduction of cats. 
Neutering should not be overused. We do not argue that Anatolian cat populations in cities could be monitored, but there is no reason to alter every living cat, especially not in less populated and rural areas of the country. If neutering campaigns aim to neuter every single cat living outside, it becomes a problem, because it is no longer a population control, it is a plan for extinction of natural cats. Neutering in massive numbers, negatively affects the gene pool of the Anatolian cats: less cats means more more inbreeding and more diseases.
Sterilization of females is more complicated and difficult than that for male cats. Unspayed males may cause stress  for the whole population - they involve into fights, which result into noise and complaints from the communities, injuries and transmission of diseases. Neutering campaigns, which want to manage the populations rather than completely eliminate them, should target males first instead of females. 
 * Face reality: cats like any other animals, unfortunately, will die. Most of kittens will not survive due to disease, congenital abnormalities, blood type incompatibilities; some will be abandoned or eaten by their mothers. While we can do our best to treat the diseases and reduce human caused suffering, it is not possible to save every cat.
All animals suffer, because this is how nature operates, on survival and death cycle. Getting rid of one species of animals, may make some people feel better about themselves, but this will not solve the problem of animal suffering. 

Human caused problems:

A frightened cat from Sur, Diyarbakır. 
Photo by Özge Özgün

Ø Wars, terrorism and with it associated damage
These tragic events destroy lives of many people, however cats are also negatively affected. They get killed, suffer from injuries, unable to access a veterinary care.The food may be scarce, and in some cases, when people face with starvation, cats may be eaten.

Anatolian cat from war torn area in Aleppo Syria. 
Photo by Maher Akraa

Ø Cats injured and killed by irresponsible drivers
Clearing cats from the streets is not a solution! Why do we blame victims, instead of dealing with a root cause? Policy-makers should focus to develop better strategies to encourage a safer driving. Irresponsible drivers who speed up their cars should receive harsh punishments, because they put other people's lives in danger. It is important not to ignore this problem because the prevention of traffic accidents would not only save the lives of cats and other animals, but many more humans. In a far future, self-driving vehicles may prevent the accidents caused by human errors.

Ø Cruelty and abuse by individuals
There are no simple solution. Cats which have no owners should have the same legal status as owned cats. Harsher punishments should be implemented for animal abusers and their crimes should be scrutinized openly. Mentally ill cat abusers should receive the treatment. 

Cats on and in garbage container. Unmanaged and easily accessible waste poses a risk of poisoning and injury.

Ø Easily accessible garbage containers 
Many cats do not wander to containers, because they feel hungry; cats are attracted to the strong smells, while disgusting for humans, they are pleasant for cats. Cats that look for food in garbage containers, may get injured and poisoned. Local municipalities should provide closed garbage containers, inaccessible for cats. 

Ø Unprotected car engines enabling cats to hide inside
Many cats and kittens are killed or severely injured when a car’s engine is started up while they are still under the hood of the car. Do not start car if you have doubts that a cat may be trapped in your engine. The permanent solution to this problem would be to block the entrance to engine openings. Car service providers should be encouraged to provide the protective shields, for example, a wire screen that would prevent cats from entering into cars. Finally, why can’t car manufacturers make it impossible for cats and other animals to crawl into the engines?

Ø Uneducated owners who fail to look after their cats
Cats are injured or killed after falling out of windows and balconies. This is known as high-rise syndrome, is not a syndrome at all but the irresponsible behaviour of humans. The owners should not allow their cats near to windows and balconies in order to prevent falls. Also they should not let their cats to access poisonous substances and plants.
Cat owners should be encouraged to seek the basic knowledge how to take care of their cats, try to understand cat behavior, health and how to achieve the well-being for their pets. Educational projects would definitely benefit welfare of cats kept as pets. 
To have any home, even a bad one, is not better for a cat; many cats will be happier living on the street and loved by their communities rather than with a person or a family who fails to meet cat's needs.Without educating people about cats and their care, the promotion of cat ownership will only result to more miserable cat's lives.

Ø Shelters. Healthy cats should not be kept in shelters. Shelters should serve needs of animals which need a special care and otherwise would die if left on their own. Majority of cats will be happier outside rather than caged in shelters or kept in overcrowded sanctuaries. Collecting healthy, capable to live outside cats equals to imprisoning them and may have serious implications on cat’s health and well-being. 
It is not possible to find a home for every cat, for this reason the resources should be used for cats which really need a home: injured, ill, blind and deaf etc. cats. 

Ø Cat hoarders are individuals who gather cats from the streets and keep them in their apartments or houses. As time goes, cat hoarder collects more and more cats. Many hoarders will start to feel so overwhelmed and unable to care for their cats anymore. A hoarder's house will be unsanitary, cats will get ill and stressed due to overcrowded environment. A hoarder may run out of money and ask for donations and help, while wealthy hoarders will continue their habits under private sanctuary or shelter. Neighbours may intervene and complain about hoarder's behavior. At the end, cats are usually taken from the hoarder, often in a poor health and psychologically damaged. 
Hoarding does not help cats, it harms them. If you suspect someone is a hoarder, contact local authorities and help to remove the cats from hoarder's home before any greater damage is done.

Ø Dogs
Although dogs are natural enemies of cats, there are examples of beautiful relationships between Anatolian cats and stray dogs. 
Some dogs may chase, attack the cats and kill the kittens. Dogs roaming in packs are especially dangerous. 
Since the majority of dogs emerged from the abandoned pets, it is a human responsibility to sterilize them.

Ø Cat breeding 

Some say, there is nothing wrong with cat breeding, as long breeders take care of their cats. Of course, cat breeders have to care for the cats they breed, otherwise this would be an animal abuse. After all they bring these cats to the world and are responsible for them.
The current Western welfare model serves the interests of cat breeders very well. First of all, the unnatural selection, caused by overuse of neutering, reduces the desired phenotypes, like certain colors and frequency of longhair gene and remove the friendliest cats from the gene pool. Second, makes it harder to find kittens, and most people will prefer to adopt a kitten rather than adult cat. 
Problems with cat breeding: Many inherited diseases in breeds resulted from unscientific choices of breeders. Breeders breed cats for some superficial trait, but at the same time, they mess up with other genes. Breeders cannot know from the look alone if a cat carries a mutation of some serious disease. Therefore, many diseases get fixed into breeds unknowingly. 
There are other ethical problems: Cat breeding normalizes disabilities (example: cats without fur or with short legs) and promotes a cat as a commodity and a product. Cat fancy is also guilty for spreading a pseudoscience as a way to promote their breeds, for example, “hypoallergenic breeds" lie. Lastly, cat breeders do absolutely nothing good for natural cats. Breeders known that natural cats will be replaced by their invented breeds, which are shown as superior and better to natural cats. Two contrasting views help to achieve this idea perfectly: breeds are portrayed positively, on the other hand, the natural cats have a very negative image. The only good reason to adopt a natural cat, as stated, is to “save its life”. Any good traits, such as attractive look, friendly personality, “charisma” of natural cats are not regarded highly, but these traits are much exaggerated in descriptions about breeds. How can we expect more people will be willing to adopt a natural cats instead of buying a cat from the breeder, when we promote breeder’s established hierarchy, that natural cats are inherently inferior to breeds?

Cat breeders are able to escape most of criticism in Turkey, because people are told that pet shops are the real problem. The fact is, most of pet shops no longer sell cats. And if some do, they are just retailers, not producers. But how come there are still so many breed cats for sale? Where people get their British and Scottish Folds? Not from petshops, but from cat breeders. Anyone who breeds cats and sells them for a profit is a breeder. Most of breeders no longer supply cats to pet shops, but sell them on internet pages and social media. 

We should not glorify breeds, do not promote them on social media, such as in Facebook, in blogs or cat related sites. If people can be convinced to stop buying breed cats, breeders will go out of business. In this case, more natural cats will be adopted. Is it not what welfare organizations want?

Final word:

Human and cat bond is ancient, older than 10,000 years. It had begun right here, in Anatolia. The descendants of ancient felines currently live almost everywhere in Anatolia, from populated cities to the most remote areas. These cats became an inseparable part of community life and Anatolian culture. Humans put a significant emotional, social and cultural importance into cats. This is why the scientific study of cats is necessary and valuable. Furthermore, because humans care about cats a lot, causing a harm and ignoring welfare of cats is morally unacceptable. The Anatolian cats are changing the way we think about cats, especially the ignored group of cats - the natural cats. This project goes against all the ideologically motivated misinformation and pseudoscience, which infested every aspect of cat related fields. But it does not mean nothing can be changed. The antidote for all that misinformation is of course, more feline science!

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25.  Driscoll C, Yamaguchi N, O’Brien SJ, Macdonald DW, 2011, A suite of genetic markers useful in assessing wildcat (Felis silvestris ssp.) - domestic cat (Felis silvestris catus) admixture. Journal of Heredity 102: S87-S90. F. s. catus and lybica share the same marker in admixture analysis for detection of  "pure" Felis silvestris (European wildcat).
26. Şahin, E. Y. (2010). Türkiye'de bulunan bazı kedi ırklarının D-Loop polimorfizminin araştırılması (Doctoral dissertation, Selçuk Üniversitesi Sağlık Bilimleri Enstitüsü).
27. Süreyye Somer,, Tekir Kedisi, 2002
28. Altunok, V., Yüksek, N., Berkman, C. C., Ağaoğlu, Z. T., & Togan, İ. (2011). Genetic Structure and Variation of Van Cats. Biochemical genetics, 49(7-8), 511-522.
29. The Angora cat standart, Ek-36 Ankara Kedisi, Resmî Gazete no: 27075, 2008, December 5, Tarım ve Köyişleri Bakanlığı. 
30.  Soysal, M. I. (2007). Native Animal Genetic Resources of Türkiye.
31.  Ishida, Y., David, V. A., Eizirik, E., Schäffer, A. A., Neelam, B. A., Roelke, M. E., ... & Menotti-Raymond, M. (2006). A homozygous single-base deletion in MLPH causes the dilute coat color phenotype in the domestic cat. Genomics, 88(6), 698-705.
32. Vella, C. M., Shelton, L. M., McGonagle, J. J., & Stanglein, T. W. (1999). Robinson's genetics for cat breeders and veterinarians (No. Ed. 4). Butterworth-Heinemann Ltd.
33.  Stelow, E. A., Bain, M. J., & Kass, P. H. (2016). The relationship between coat color and aggressive behaviors in the domestic cat. Journal of applied animal welfare science, 19(1), 1-15.
34.  DK Publishing (Dorling Kindersley), The Cat encyclopedia the definitive visual guide (2014), page 64-65
35.  Gandolfi, B., & Alhaddad, H. (2015). Investigation of inherited diseases in cats Genetic and genomic strategies over three decades. Journal of feline medicine and surgery, 17(5), 405-415.
36.  Lyons, Kurushima, 2012, A Short Natural History of the Cat and Its Relationship with Humans, Chapter 42, page 1254-1262. Susan Little, (ed), The Cat: Clinical Medicine and Management.
37.  Cat Ancestry - Tracing the Lineage of Your Feline, University of California at Davis
38.  Yilmaz, O., Akbag, H. I., Coskun, F., Ozcetin, T., & Ertugrul, M. (2014). Çanakkale İlinde Yetiştirilen Kısa Tüylü Türk Kedisi’nin Tanimlanması. Türk Tarım ve Doğa Bilimleri, 7(7), 1993-1997.
40.  Hébert, J. M., Rosenquist, T., Götz, J., & Martin, G. R. (1994). FGF5 as a regulator of the hair growth cycle: evidence from targeted and spontaneous mutations. Cell, 78(6), 1017-1025.
41.  Higgins, C. A., Petukhova, L., Harel, S., Ho, Y. Y., Drill, E., Shapiro, L., ... & Christiano, A. M. (2014). FGF5 is a crucial regulator of hair length in humans. Proceedings of the National Academy of Sciences, 111(29), 10648-10653.
42.  Lyons, L. A. (2015). DNA mutations of the cat. The good, the bad and the ugly. Journal of feline medicine and surgery, 17(3), 203-219.
43.  Godfrey, Genetic Welfare Problems of Companion Animals, Persian – Dermatophytosis (2011)
44.  Drögemüller, C., Rüfenacht, S., Wichert, B., & Leeb, T. (2007). Mutations within the FGF5 gene are associated with hair length in cats. Animal genetics, 38(3), 218-221.
45.  Housley, D. J. E., & Venta, P. J. (2006). The long and the short of it: evidence that FGF5 is a major determinant of canine ‘hair’‐itability. Animal genetics, 37(4), 309-315.
46.  Dierks, C., Mömke, S., Philipp, U., & Distl, O. (2013). Allelic heterogeneity of FGF5 mutations causes the long‐hair phenotype in dogs. Animal genetics, 44(4), 425-431.
47.  Kehler, J. S., David, V. A., Schäffer, A. A., Bajema, K., Eizirik, E., Ryugo, D. K., ... & Menotti-Raymond, M. (2007). Four independent mutations in the feline fibroblast growth factor 5 gene determine the long-haired phenotype in domestic cats. Journal of Heredity, 98(6), 555-566.
48.  Bach, L. H. (2010). Analysis of FGF5 and construction of a high-resolution radiation hybrid panel for the domestic cat. University of California, Davis.
      90 % of all longhaired pedigreed breeds and many natural cats had this particular mutation- c.475A>C (found in 22 out of 24 populations).
49.  Kienzle E, Moik K., 2011, A pilot study of the body weight of pure-bred client-owned adult cats, Br J Nutr. 2011 Oct;106 Suppl 1:S113-5. doi: 10.1017/S0007114511001802. 
     "In all breeds, there was a marked sexual dimorphism with heavier males than females. Even though this was more marked in some breeds than others, the difference is too big to allow a calculation of the mean of a group of cats regardless of their sex" This observation applies to all cats, not only to selected breeds.
50.  Benjamin S. Arbuckle, 2012, A Companion to the Archaeology of the Ancient Near East, part 11. Animals in the Ancient World p. 210, editor: D. T. Potts, Wiley-Blackwell.
51.  Yamaguchi, N., Kitchener, A., Driscoll, C. & Nussberger, B. 2015. Felis silvestris. The IUCN Red List of Threatened Species 2015: e.T60354712A50652361
52.  For example, a veterinarian Erkan Morgül in one video site describes the Norwegian Forrest cat and shows a longhaired Anatolian cat as example of the “breed”.
53.  Calling the Anatolian cats as breeds is very common in social media and pet related sites. For example, a number of people identified their cats as breeds, such as Bombay, Maine Coon, Russian Blue etc., none of which are exist in Turkey.  while other site offered a search for adoptions based on cat’s breed.  Of course, many cats were given random breed labels to increase chances for adoption (obvious examples are Japanese Bobtail, Burmese, Sphynx, Brazilian Shorthair (?) etc.). But some, especially if cats got certain colors of fur slightly resembling those observed in breeds, were labeled as breeds not accidentally: tabbies were Bengals, black cats – Bombay, longhair - Norwegians etc.
54.  Starkovich BM, Stiner MC. 2009. Hallan Çemi Tepesi: High-ranked game exploitation alongside intensive seed processing at the Epipaleolithic-Neolithic transition in Southeastern Turkey. Anthropozoologica 44(1): 41-61.
55.  Lentacker, A., & Cupere, B. D. (1994). Domestication of the cat and reflexions on the scarcity of finds in archaeological contexts. Colloques d'Histoire des Connaissances Zoologiques (Belgium).
56. S. T. Özçetin, 2007, Ankara kedilerinde diş yapi, tüy, büyüme, gelişme ve üreme özellikleri üzerine araştirmalar, Ph.D thesis, Ankara University, Graduate School of Natural and Applied Sciences Department of Animal Science (Özçetin claims that Anatolian cats have rounder heads than Egyptian cats based on artifacts with exaggerated features) 
57. Ottoni, C., Van Neer, W., De Cupere, B., Balasescu, A., Benecke, N., Boivin, N., Buitenhuis, H., Chahoud, J., Manaseryan, N., Monchot, H., Morales, A., Onar, V., Osypińska, M., Peters, J., Prendergast, M., Putelat, O., Spassov, N., Studer, J., Wierer, U., Decorte, R., Grange, T., Geigl, E. (2015). Ancient DNA from cats - a paleogenetics perspective into past distributions and ancient human mediated translocations of Felis silvestris. ASWA. Groningen (The Netherlands), 10-13 June 2015.
58. Russell, N. & Martin, L. (2005) The Çatalhöyük mammal remains. In hodder, I. (Ed.) Inhabiting Çatalhöyük: Reports from the 1995-1999 Seasons. Cambridge, McDonald Institute for Archaeological Research.
59. Starkovich BM, Stiner MC. 2009. Hallan Çemi Tepesi: High-ranked game exploitation alongside intensive seed processing at the Epipaleolithic-Neolithic transition in Southeastern Turkey. Anthropozoologica 44(1): 41-61.
60. Voigt, M. (2007). The splendour of women: late Neolithic images from Central Anatolia. Material beginnings: a global prehistory of figurative representation, 151-169.
61. Jenkins, E. L., 2012. Mice, scats and burials: unusual concentrations of microfauna found in human burials at the Neolithic site of Catalhoyuk, Central Anatolia. Journal of Social Archaeology, 12 (3), 380 - 403.
62. Russell, N., & Martin, L. (1995). The Çatalhöyük mammal remains. Inhabiting Çatalhöyük: reports from the, 1999, 33-98.
63. Spotte, S. (2014). Free-ranging cats: Behavior, ecology, management. John Wiley & Sons.
This is a good example. The author of this book displays very negative biases towards natural cats;
the references he uses to support his views are mainly taken from the scientific literature.
64.Riesen, M. (2007). The Pet Overpopulation Crisis: How Training the Public Can Make a Difference. JACAB, 1(1), 22-27.
65. Bartlett, P. C., Bartlett, A., Walshaw, S., & Halstead, S. (2005). Rates of euthanasia and adoption for dogs and cats in Michigan animal shelters. Journal of Applied Animal Welfare Science, 8(2), 97-104.
66. Denny, E. A., & Dickman, C. R. (2010). Review of cat ecology and management strategies in Australia. Invasive Animals Cooperative Research Centre, Canberra.
67. Farnworth, M. J., Dye, N. G., & Keown, N. (2010). The legal status of cats in New Zealand: A perspective on the welfare of companion, stray, and feral domestic cats (Felis catus).
68. Podberscek, A. L. (2009). Good to pet and eat: The keeping and consuming of dogs and cats in South Korea. Journal of Social Issues, 65(3), 615-632.
69. Czaja, R., Wills, A., Hanitriniaina, S., Reuter, K. E., & Sewall, B. J. (2015). Consumption of Domestic Cat in Madagascar: Frequency, Purpose, and Health Implications. Anthrozoös, 28(3), 469-482.
70. Gürler, A. M., Meli̇koğlu, B., & Osmanağaoğlu, Ş. (2011). A Historical evaluation of animal protection efforts of non-governmental organizations in Turkey. Kafkas Üniversitesi Veteriner Fakültesi Dergisi, 17(6), 901-908.
71. Chartrand, T. L., Fitzsimons, G. M., & Fitzsimons, G. J. (2008). Automatic effects of anthropomorphized objects on behavior. Social Cognition, 26(2), 198.
72. Epley, N., Waytz, A., Akalis, S., & Cacioppo, J. T. (2008). When we need a human: Motivational determinants of anthropomorphism. Social cognition, 26(2), 143-155.
73. Neilson, J. (2004). Thinking outside the box: feline elimination. Journal of Feline Medicine and Surgery, 6(1), 5-11.
74. Cevizci, S., & Bakar, C. (2013). Halk Sagligi bakisiyla Toxoplasma gondii. Turkish Journal of Public Health, 11(1), 45.
The authors of this paper repeat many times these phrases: "sokağa bırakılan" (left on the street), "Sokağa terk edilen hayvan" (an animal abandoned on street) as it was an obvious fact. Of course, where they get these "facts" from is a mystery. The authors fail to acknowledge that most of cats come from feral, freely living populations which do not have owners, other than being looked after by the communities. Surely, the solution to prevent Toxoplasma gondii from spreading do not require much thinking: Just get rid of all cats living on the streets, as the Western welfare model dictates.
75. The tendency to image that cats living outside have a desire to "have a home" is evident in many publications. For example, an exercise book published by Ministry of national Education "Hayat Bilgisi - 1 Ders ve Öğrenci Çalışma Kitabı" (2014-2015) shows a conversation between two children who feel sorry for cats walking in the rain: "Every living thing needs a home. The mother cat and her kittens are getting cold because they do not have a home". Another responds: "Let's take a permission from our mother. Let's make a house for them and give some milk".
76. "İstanbul’da kedi, dünyada örneği olmayan bir şey" interview with Ceyda Torun by İpek İzci; 02/21/2016; Hürriyet
77. Montague et al., 2014, Comparative analysis of the domestic cat genome reveals genetic signatures underlying feline biology and domestication, Proceedings of the National Academy of Sciences of the United States of America.
78. H. O. Heyne et al., 2014, Genetic Influences on Brain Gene Expression in Rats Selected for Tameness and Aggression, Genetics 04/2014; DOI: 10.1534/genetics.114.168948
79.  Seitz, P.F.D. (1959). Infantile experience and adult behavior in animal subjects. II. Age of separation from the mother and adult behavior in the cat. Psychosomatic Medicine, 21, 353–378. 
80. Lowe, S.E. & Bradshaw, J.W.S. (2001). Ontogeny of individuality in the domestic cat in the home environment. Animal Behaviour, 61, 231–237. 
81. McCune, S. (1995). The impact of paternity and early socialisation on the development of cats’ behavior to people and novel objects. Applied Animal Behaviour Science, 45, 109–124. 
82. Reisner, I.R., Houpt, K.A., Erb, H.N., et al. (1994). Friendliness to humans and defensive aggression in cats: the influence of handling and paternity. Physiology and Behavior, 55, 1119–1124. 
83. Feuerstein, N. & Terkel, J. (2008). Interrelationships of dogs (Canis familiaris) and cats (Felis catus L.) living under the same roof. Applied Animal Behaviour Science, 113, 150–165. 
84. Kuo, Z.Y. (1930). The genesis of the cat’s response to the rat. Journal of Comparative Psychology, 11, 1–35. 
85. Caro, T.M. (1980a). The effects of experience on the predatory patterns of cats. Behavioral and Neural Biology, 29, 29–51. 
86. Wyrwicka, W. (1978). Imitation of mother’s inappropriate food preference in weanling kittens. Pavlovian Journal of Biological Science, 13, 55–72. 
87. Reisner, I.R., Houpt, K.A., Erb, H.N., et al. (1994). Friendliness to humans and defensive aggression in cats: the influence of handling and paternity. Physiology and Behavior, 55, 1119–1124. 
88. Turner, D.C., Feaver, J., Mendl, M., et al. (1986). Variation in domestic cat behaviour towards humans: a paternal effect. Animal Behaviour, 34(6), 1890–1892. 
89. Cieslak M, Reissmann M, Hofreiter M, Ludwig A. (2011). Colours of domestication, Biol Rev Camb Philos Soc. Nov; 86(4):885-99. doi: 10.1111/j.1469-185X.2011.00177.x.
90. Fred W. Allendorf, Conservation and the genetics of populations (2012), Wiley. "As deleterious mutations accumulate, population size may decrease further and thereby accelerate the rate of accumulation of deleterious mutations" (p. 290). "We must first maintain healthy habitats and large wild populations, because only in large populations can natural selection proceed efficiently" (p. 320) 
91. Allendorf, F. W., & Hard, J. J. (2009). Human-induced evolution caused by unnatural selection through harvest of wild animals. Proceedings of the National Academy of Sciences, 106 (Supplement 1), 9987-9994.
92. Stenseth, N. C., & Dunlop, E. S. (2009). Evolution: unnatural selection. Nature, 457(7231), 803-804.
93. Cameron-Beaumont, C., Lowe, S. E., & Bradshaw, J. W. S., 2002, Evidence suggesting preadaptation to domestication throughout the small Felidae. Biological Journal of the Linnean Society, 75 (3), 361 - 366. 10.1046/j.1095-8312.2002.00028.x 
94.Sunquist, F., & Sunquist, M. (2014). The Wild Cat Book: Everything you ever wanted to know about cats. University of Chicago Press. (pages: 205-244)
95. Stanton, L. A., Sullivan, M. S., & Fazio, J. M. (2015). A standardized ethogram for the felidae: A tool for behavioral researchers. Applied Animal Behaviour Science, 173, 3-16.
96. Collard, R. R. (1967). Fear of strangers and play behavior in kittens with varied social experience. Child Development, 877-891.
97. Frank, D., & Dehasse, J. (2004). Differential diagnosis and management of human-directed aggression in cats. Clinical techniques in small animal practice, 19(4), 225-232.
98. Woolliams, J. (2012). Influence of genetics and inbreeding on disease. In Practice, 34(4), 196.
99. Todd, N. B., & Todd, L. M. (1976). Mutant allele frequencies in the domestic cats of Turkey and Greece. Genetica, 46(2), 183-192.
100. Roy Robbinson, 1972 Mutant Gene Frequencies in Cats of Cyprus, Theoretical and Applied Genetics 42, 293- 296. 
101. M. Ruiz-Garcia, 1994, Genetic profiles from coat genes of natural Balearic cat populations: an eastern Mediterranean and North-African origin, Genet Sel Evol.; 26(1): 39–64 
102.102. Gould, D. (2011). Feline Herpesvirus-1 Ocular Manifestations, Diagnosis And Treatment Options. Journal Of Feline Medicine And Surgery, 13(5), 333-346.
103. Stiles, J., & Kimmitt, B. (2016). Eye examination in the cat Step-by-step approach and common findings. Journal of Feline Medicine and Surgery, 18(9), 702-711.
104. Maes, R. (2012). Felid herpesvirus type 1 infection in cats: a natural host model for alphaherpesvirus pathogenesis. ISRN veterinary science, 2012.

Note: the list of sources is not complete - will be updated soon.

Authors: Perla Aksoy, Batu Aksoy 
Publication date: 8/18/2016

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