What is the Anatolian Cat? To answer this question we have to delve deeper into the history and the genetics of cats.
In the first part of the Anatolian Cat (Anadolu Kedisi) series, we will investigate the complex, yet fascinating, origins of cats.
Which wildcats were the ancestors of Anatolian cats and other domestic cats? Were cats really “domesticated” in Egypt?
When, where and how cats began their unique relationship with humans?
Domestic cat and its relatives
Before exploring the history of domestic cat, we should know how the domestic cat is related to other cat species: we need to look at the taxonomy of the domestic cat.
Pseudaelurus - the ancestor of cat family Felidae. Courtesy: Columbia University Press.
All 39 living cat species share a last common ancestor known as Pseudaelurus, a cat that lived about 10-15 million years ago (1).
Figure 1. Feline family tree (Felidae). The domestic cat belongs to genus Felis, wildcat species Felis lybica and subspecies F. lybica lybica. Image courtesy: Anadolu Kedisi
Cat species can be categorized into 8 separate lineages called genus (Figure 1). When it comes to the domestic cat, its genus is Felis. Felis genus appeared about 3.36 million years ago (2). Sand cat, jungle cat, black-footed cat, European wildcat, and the closest domestic cat relatives, African and Asian wildcats - all are a part of Felis genus.
The domestic cat belongs to the wildcat species called Felis lybica – the cats that historically inhabited areas in the Near East, Africa, and Asia. Three subspecies of Felis lybica have been identified so far: the Asiatic wildcat (Felis lybica ornata), South African wildcat (Felis lybica cafra) and the Near Eastern wildcat (Felis lybica lybica) (3). The domestic cat is descended from the Near Eastern wildcat, therefore, the domestic cat’s taxonomic name should be Felis lybica lybica (4, 5).
However, taxonomists choose to ignore the genetic and archeological data that supports the classification of domestic cat as Felis lybica lybica species, and instead, they call it as separate species “Felis catus” (6).
Felis domesticus, the name used to describe a domestic cat in older publications, is invalid and not used anymore (7).
Photo credit: Michael Lindle
A note on why we use the term “taming” instead of “domestication”
We describe cats from ancient times, or to be exact, the wildcats that lived with humans, throughout this article as “tamed”. This may sound like an unusual choice knowing that lots of publications prefer the term "domestication".
The central point of the first part of the article is the very beginnings of cat and human relationship in the Neolithic period - we rarely go beyond that. Therefore it is not relevant to examine whether humans successfully domesticated cats or not.
There is a little doubt that cats at that time would still be considered to be wildcats. Since there was no outgoing direct human selection, and wildcats living with humans were not isolated from their wild population, taming seems to be a more appropriate term than domestication.
Additionally, the “domestication” does not have a single agreed definition (8, 9). Scientists still debate whether the domestic cat should be categorized as domesticated or not (10,11,12). So we are going to explore the domestication topic separately in the second part.
Sardes Artemis temple Salihli Manisa. Photo credit: Özkan Aras
Where were cats first tamed?
For a long time, it was believed that domestic cats originated and were “domesticated” in Egypt 4000 years ago, and from there spread to other parts of the world (13). It is not difficult to understand why this belief persisted. Cats were a part of the well documented religious cult. Ancient Egyptians left many artifacts portraying domestic cats along with 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” the cats, or were they?
The current discoveries in archeology and genetics are already rewriting the history of the domestic cat. The evidence suggests that the importance of Egypt in cat “domestication” has been greatly overstated.
No sufficient evidence for cat “domestication” in Egypt or China
Figure 2. Egyptian cat mummies and the specimens from the site of Quanhucun in China that belong to leopard species.
Despite its popularity, the hypothesis that ancient Egyptians “domesticated” cats lack support from science.
The study on 2 and 3 thousand-year-old Egyptian mummies revealed that mummified cats actually were very similar to the current populations of Egyptian cats. It appears that the domestic cat arrived in Egypt after it had already been tamed elsewhere (14). Although it was commonly accepted that cats started to live with humans in Egypt 4000 years ago, the discovery of cat burial from ancient Egyptian town Hierakonpolis predates this relationship by nearly 2000 years (15).
According to another study, cats from unidentified species appeared in China in the 5,300-year-old agricultural village of Quanhucun. These cats fed on rodents that ate stored grains (16). Some sources wrongly interpreted this study as an evidence, that cats were domesticated in China (17). The study described the commensal relationship between humans and wild cats which may precede the domestication (18). Commensalism is a relationship between two species of organisms sharing the same environment when one benefits from that relationship without harming the other. Quanhucun cats clearly were not domesticated but the conditions (the abundance of rodents in a village, living in close proximity to humans) could initiate the taming of those cats (19).
The study may not be relevant anymore in tracing the origins of the domestic cat because the cats from ancient Chinese village were identified to belong to leopard cat species (Prionailurus bengalensis) (20).
Why cats decided to live with humans
Photo credit: Bülent Kotaman
Anatolia was a home for the earliest permanent settlements and a place, where the agriculture started (21). Many animals and plants were domesticated at that time (22).
The most important crop, einkorn wheat was also domesticated in Southern Anatolia, Karaca Dağ Mountain (23, 43), close to the location of Göbekli Tepe. Göbekli Tepe is a 12,000-year-old temple which gives the earliest evidence of the first organized human societies and marks the transition from hunting and gathering to farming (24).
Soon after the first villages appeared in Anatolia, and humans started practicing agriculture, ancestors of house mice (Mus musculus), and wildcats were drawn to these villages (25).
Throughout most of the history, cats have had a commensal relationship with humans: cats hunted rodents and fed on waste and humans tolerated their presence (10, 21).
However, cats were not only tolerated but could also be valued as human companions too. The oldest known finding that provides evidence of a close relationship between a human and a cat, is ~9,500 years old cat and human burial from Cyprus (26). No cats were present in Cyprus until early farmers from Anatolia brought the wildcats to the island by boat (27). These wildcats had been tamed, meaning that the relationship between cats and humans in Anatolia dates back at least 10,000 years.
Photo credit: Sabri Keleş
It is very unlikely that Neolithic villagers attempted to domesticate the wildcats roaming in their settlements. However, the vulnerability of the wildcat could be a starting point of the cat-human relationship.
Imagine the situation, when humans decided to respond to a meowing of a kitten, left by its mother, and began to look after it. The cuteness and helplessness of the kitten triggered the nurturing feelings in humans (28, 29). Those kittens had an opportunity to interact with humans early on, and if this interaction continued, the taming of wildcats became inevitable (30).
Photo credit: Ogün Çıtlak
However, the tendency to tameness is not exceptional to the domestic cat or F. lybica wildcats but was observed in many cat species, including the distant relatives of domestic cats: caracal, lynx, and leopard (31). The finding from China (16) demonstrates that leopards (Prionailurus bengalensis) had a very similar relationship with humans (19), as did Anatolian farmers with F. s. lybica wildcats at first. However, whatever the reason, human and leopard relationship did not last.
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 waste left by humans and hunted small animals, which lived in human settlements. Some cats were probably even fed by humans. This allowed the tame cat population to increase and dominate over the untamed wildcats in the areas they inhabited, although wild and tame populations never became separate.
Finding the ancient Anatolian Cat
Figure 3. Illustration of the 9,500-year-old cat and human burial from Shillourokambos site, Cyprus.
Figure 3a. Photo showing the burial of the cat (lower skeleton) and the human (above) at Shillourokambos.
Human and cat burial in Cyprus (26) is an important discovery because it points towards the Anatolian origin of the domestic cat (Figure 3 ans 3a).
Unfortunately, archeologists have not yet located any similar burials from Anatolia. Cat bone fragments were found in ancient settlements, such as Çatalhöyük (32, 33, 34), Hallan Çemi (35), Aşıklı Höyük (4), Bademağacı and other sites. But it cannot be known if cats found in these settlements were tame or wild.
Anatolian cat walking at the top of beehive houses in Harran, Şanlıurfa. Photo credit: Howard Koons .
Research shows that inhabitants of one of the largest Neolithic settlement Çatalhöyük (Konya province), welcomed the small predators and wildcats into their houses because these animals helped to deal with mice invasion (33, 36).
A recreation of Çatalhöyük.
Yet, randomly distributed remains of cats in Çatalhöyük and in other settlements, do not give us any insights into the cat and human connection. It is possible that some wildcats were human companions, yet no documented evidence for their relationship survived. The remains of cats or even their burials could be in the area outside the settlements, often missed by archeologists (37).
Feline symbolism frequently appears in the Anatolian art from the Neolithic period (38). Female figurines from Çatalhöyük and Hacılar are accompanied by felines, probably leopards or lions. It is possible, that some of these figurines and other artifacts (Figure 5) depict the domestic cat ancestors, as seen in the 7,500-year-old Hacılar example, where a woman holds an animal that looks similar to the domestic cat (Figure 4).
Figure 4. Mother Goddess holding a small feline; Hacılar 5,500 BC. Photograph by Özkan Aras, drawing by James Mallart
Another example is a plaster head from Çatalhöyük, dated circa 8,500 years (Figure 5, left). Archeologist and a director of Çatalhöyük Research Project, Prof. Ian Hodder thinks that this plaster head portrays a feline (39).
But probably the most remarkable finding is a cat head of carved stone from Shillourokambos site, southern Cyprus (40). This sculpture is almost 10,000 years old (Figure 5, right).
Figure 5. Artifacts that possibly depict the heads of cats from Çatalhöyük and Shillourokambos sites.
Figure 5a. Cat head shaped ritual vessel, 1600-1700 BC (Alişar Höyük, Central Anatolia)
Research confirms that Anatolia is likely a homeland of cats
Archeological evidence alone is not sufficient to determine where the domestic cat was first tamed. The genetic studies of recent cat populations already confirmed that only one species of wildcat gave rise to all the domestic cats. And it has been already confirmed that this wildcat lived in the area vaguely defined as the “Near East” (5).
Anatolian cat at the library of Celsius, Ephesus, Turkey. Photo credit: Salih Altuntaş
To reveal the exact location of where domestic cat ancestors came from, we need to study the ancient DNA.
What is the ancient DNA? It is DNA extracted from an ancient material, such as bones of felines discovered in ancient settlements.
The study (Ottoni et al., 2017) extracted Mitochondrial DNA from remains of cats found in Neolithic settlements like Aşıklı Höyük, Bademağaçı, and younger sites, Sagalassos, Demirci Höyük. The oldest specimen of the cat in this study was almost 10,000 years old and came from Aşıklı Höyük site. The primary analyses showed that cats from these places belonged to F. l. lybica species (4). It may come as a surprise for many, who believed that F. l. lybica wildcats could not be found in Anatolia. It turns out that F. l. lybica wildcats lived in Anatolia long before the arrival of farmers, and its range extended up to Balkans and Caucasus.
Figure 6. Felis lybica lybica (former name: Felis silvestris lybica) maternal lineages from which the domestic cat originated: “Clade A” is Anatolian, Clade B - Anatolian & Levantine, Clade C origin is open to question. Courtesy: Nature Ecology & Evolution
According to a study, the earliest steps of cat taming were really taken in Anatolia.
F. lybica wildcats’ maternal lineages, the most ancient lineages, classified as Clade A and B, originated from the Anatolia Region (Figure 6).
Clade C appears to be acquired more recently - about 8th century BC. This lineage is associated with Greek and Roman expansion. Researchers speculate that Clade C came from Egypt, however, the event of mass importation of cats from Egypt to Anatolia is highly unlikely. It is plausible that this lineage was present in Anatolia before, and could be detected in samples of cats from South Eastern Anatolia and Northern Syria (samples from these regions were not included in a study). The data is limited, and more samples and advanced genomic analyses are necessary.
Anatolian cat in the ruins of the ancient city of Palmyra, Syria. Photo credit: Craig Jenkins
Apparently, cats from Anatolia did migrate to Egypt and beyond: Clade A can be found in cats throughout Africa.
Researchers think that other minor lineages of F. l. lybica (Clade D and E) have been incorporated more recently through admixture with wild populations. Surprisingly, Asian wildcat, F. l. ornata, also managed to get into the domestic cat genetic history (4, 41). Research suggests that Asian wildcat was brought via trade routes to the Near East and Europe and interbred with local wildcats (4).
The research on domestic cat origins still continues and may yield to some unexpected findings. The research project led by archaeozoologist Wim Van Neer is currently in progress. Because researchers will sequence whole genomes of ancient and modern cats, the results are expected to be more complete and reliable. The study plans also to include 10 cat samples from Çatalhöyük site (42). The study may reveal the origin of Clade C and expand our current knowledge of the domestic cat history.
Photo credit: Carlos Vera
“Neolithic revolution” brought profound changes in both human and cat history. The beginning of agriculture and animal domestication led to the emergence of sedentary communities that later grew into the first farming villages. These changes created an opportunity for the development of human and cat relationship which began when F.l. lybica wildcats started hanging around the farming communities in Anatolia, about 10,000 years ago.
Throughout these thousands of years, humans had no intention of changing a cat in any way. Although cats still genetically resemble their wild ancestors, many still believe that cats are domesticated.
In the second part of the series, we will talk about cat domestication: is a cat really domesticated or just tame?
Author: Feline Scientist
Cover photo: Ekrem Güngör
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