Cats are one of the most popular pets in many Western countries. Despite our close relationship with these animals, we still have very little knowledge of their origins.
Two important discoveries gave us some clues where cats possibly came from 9,500-year-old cat and human burial from Cyprus suggested that wildcats were first tamed in Anatolia (1), and a study conducted in 2007 confirmed that all cats originated from a few Felis silvestris lybica wildcat lineages somewhere in the Near East (2).
However, the hypothesis that cats were domesticated in ancient Egypt from local wildcats and from there spread to other parts of the world, has always been more popular among the general public and scientists alike.
So where is exactly the domestic cat’s homeland? Is it somewhere in the Near East or in Egypt? Did humans domesticate cats more than once? A widely publicized study published in Nature Ecology & Evolution journal attempted to answer these questions.
The study is valuable but is far from being groundbreaking. It has been making headlines in many media outlets.
We have reviewed numerous media reports, some of which we used as examples (3-21). As we will see soon, the mainstream media picked up on Egyptian and Viking narrative enthusiastically. Here is the summary:
♦ Cats were domesticated twice (“two waves”): 10,000 years ago by first farmers, and later by ancient Egyptians. Many sources do not name Anatolia, only the Middle/Near East, southwest Asia or Fertile Crescent (5, 6, 7, 8, 9, 11, 13, 16). A video from Nature ignored the name of Anatolia Region completely, only mentioning the “Fertile Crescent” (22). It is said that both Anatolian and Egyptian wildcats supposedly contributed to the gene pool of the domestic cat. “Today’s cats are likely a blend of both Turkish and Egyptian cats (4).”
♦ Egyptian cats were more social and tame, this is why they were more popular than Anatolian cats (4, 6, 14, 15, 16). Anthrozoologist and animal behaviorist John Bradshaw, adds that Egyptian cats were “more fun to be with but just as good at hunting mice” (18).
♦ Vikings carried cats of Egyptian origin in their ships for pest control (9, 11, 13). Vikings also supposedly helped to spread these cats all over Europe (5) and maybe “took cats with them when they made early voyages to the Americas in the 10th century CE, 500 years before Columbus (18).” It is worth to mention that Viking and cat story-line appears a second time in a connection to this study. “Viking cats” story caught everyone’s attention, even before the manuscript was submitted (24). When the manuscript of the study finally came out, nobody seemed to care about it (23).
♦ Only a few news sources included researcher’s doubts about Egyptian cat’s uncertain origin. The possibility that Egyptian cats were descendants of the Near Eastern cats, cannot be ruled out (3, 4, 10, 19).
♦ Only one source, Science magazine, very briefly questioned the hypothesis whether cats were really domesticated twice (4).
♦ Some sources included cat mummies, artifacts and similar images of ancient Egyptian cats (4, 6, 7, 10, 15, 17, 18, 20). These images were used to give emphasis on Egyptian role in a cat domestication and attract the attention of readers.
Figure 1. Who cares about Anatolia, when we have Fertile Crescent?
♦ The video from Nature journal used the outdated Fertile Crescent map and ignored the whole of Anatolia (figure 1), although the study explicitly states that it detected the ancient wildcat lineage of Anatolian origin. Moreover, the majority of cat samples of this lineage came from the Western Anatolia. Obviously, this part of Anatolia was not pictured on a map (22).
When comes to Turkish media, one misleading and a low-quality article coming from Seeker site (20), dominated. This was because the archaeology website called Arkeofili was the first one to translate the article on this topic (21), and other Turkish media outlets just copied from there.
To get a better impression how bad the article is, here is a quote: “Domestic cats over the years have bred with local wildcats, resulting in everything from the now-extinct Mexican hairless cat to the Siamese, which derived from a feline native to Thailand (20).”
The author confuses wildcat lineages with human-made cat breeds. And what is this Mexican hairless cat? The article also misleads its readers about tabby cats saying that tabby cats (without specifying it was not any tabby, but a blotched variety) appeared in Ottoman Empire as if it is another breed of cat.
The majority of claims presented in mainstream media were drawn from researchers conclusions quite accurately, but not without exaggerations.
Many people will learn about this study from the secondary sources as ones described above, not by reading the original paper. Even if media reports seem to be in line with researchers conclusions, the information is still incomplete: if we have no opinion on this study, we will take the media’s claims at face value. If we seek to understand the study better, should know what the scientists really said in their research, and look for inconsistencies in it.
Note: We refer The Near Eastern wildcats as Felis silvestris lybica in this article because this is what the study used and also to avoid the confusion that can come with using the new taxonomic name. Taxonomists renamed Felis silvestris lybica to Felis lybica lybica in 2017 (41).
Our role in this study
We have not been involved in this study directly, but we were glad to take a place of volunteer “peer review”. We discovered an abstract of the pending study before the study itself was completed. Our first contact with Dr. Claudio Ottoni was in 2015 (25). We have been following the progress of his research since then, occasionally having discussions with Dr. Ottoni.
Although the manuscript of this study was available on biology pre-print site bioRxiv since 2016, surprisingly, it received virtually no attention from the media (23). “Viking cats” story featured on a poster from a conference turned out a distraction, driving away from the original paper (24).
Once the manuscript came out, we had reviewed it and sent our observations, suggestions, and corrections, hoping that will make the final publication better. Most of the points presented in this article were discussed with the author of the study.
We have been informed months before in which scientific journal the study will be published.
Why should we care about this study?
♦ It is the first study on cats that used the ancient DNA. Ancient DNA is much more accurate and informative compared to DNA from modern populations in studying the history and origins of the domestic cat.Thanks to the ancient DNA from various locations, we can make a better hypothesis where cats were tamed, which wildcat lineages shaped the domestic cat genetics, and how populations interacted and changed over time.
♦ Unlike previous studies, this particular research has shown the significance of Anatolia in cat domestication. It confirms that the most ancient lineages of domestic cat ancestors are from Anatolia.
♦ It shows that domestic cats are no different from their wild ancestors genetically.
♦ It improves our understanding of s. lybica wildcat distribution. Before F.s. lybica was associated with North Africa, and it was assumed these wildcats are not found in Anatolia. However, this study revealed that lybica was native to Anatolia Region and that it existed there long before humans began to live in communities and invented farming.
♦ As for our Anatolian Cat project, we are pleased that a scientific publication mentions the Anatolian cat name for the first time and recognizes its existence.
What does the study claim?
It is well known that the domestic cat ancestor is a wildcat called Felis silvestris lybica*. The previous study from 2007, found that the domestic cat is a combination of a few separate lybica lineages (2). It is essential to understand that throughout the study and this article we talk about lybica wildcat lineages, not about any specific domestic cat populations, and certainly not about cat breeds.
domestic cat is classified as separate species Felis catus by taxonomists, but this research makes no distinction between wildcats and domestic cats. The domestic cat is not only too similar to its wild ancestors genetically, but also morphologically and behaviorally. Coat color remains one of the few ways we can recognize the domestic cat from the wildcat.
In this study, researchers extracted DNA from bone, teeth, skin and hair samples of 209 ancient cats. They used mtDNA (mitochondrial DNA, maternal line) to find the differences and categorize the cats.
Claudio Ottoni sampling tissue of cat mummies in the Natural History Museum, London.
The sequences of ancient and modern s. lybica wildcats were matched to the set of mitotypes/haplotypes. Mitotypes/haplotypes are varieties of mitochondrial DNA, inherited from the female lineage, so this is just a part of the story.
The study says, that 3 clades of s. lybica wildcats called IV-A, IV-B, and IV-C contributed the most to the domestic cat genetics. These lineages are regarded as ancient, while other mitotypes IV-D and IV-E were incorporated more recently from mating with wild/untamed cat populations.
The image below illustrates the wildcat lineages which made into the gene pool of the domestic cat (Figure 2).
Figure 2. Tree of mtDNA lineages: A, B and C clades of Felis silvestris lybica contributed to the gene pool of domestic cats; minor influence comes from D and E.
What this all means? After the identification of wildcat lineages, we can pinpoint their origins. Then we can find where these particular wildcat lineages come from. Finally, we move to the final step: decide from the data we have when and where domestication* of cats took place.
Researchers gave little attention to lineages IV-D and IV-E because they believed these are probably from the recent admixture between domestic and untamed wildcats. There are other lybica lineages they detected in some African cats, but these lineages were not present in cats they tested. So we will focus on IV-A, IV-B, and IV-C only.
The lineage of IV-A was found in almost 10,000-year-old site Aşıklı höyük (Central Anatolia) and it appeared in all Western Anatolian samples from the Neolithic period. Data shows that these wildcats were brought to Europe, likely during the time when Anatolian farmers spread north, therefore, the lineage IV-A was also found in cats from Bulgaria and Romania.
IV-B is present in Syria and Levant, and very likely in Southeastern Anatolia, so we can name it as Anatolian/Levantine.
IV-C lineage history is controversial. This lineage appears in the Greco-Roman period and spreads in many places of Anatolia and at the same time is transported into Europe.
IV-C lineage will turn out the most important and most cited part of the study, because of the emphasis it was given.
“The second wave” came from Egypt or somewhere else?
The study clearly states that there were two domestication events: first in Neolithic Anatolia and the second in Egypt. This is what the mainstream media reported too.
These conclusions are based upon data summarized in this, at first sight, complicated table of haplotypes (Figure 3).
Darker yellow IV-A, pale yellow IV- B, orange IV-C, light blue – IV E and D.
Novel silvestris lybica lineages are in pink color, not detected in domestic cats so far. Green – European wildcat, purple – Asian wildcat F. s. ornata.
Figure 3. The representation of cat haplotypes illustrated in colorful shapes and sorted according to the geographic origin and chronological period.
As we can see there are a lot of gaps in a table (missing data). DNA samples from many ancient Egyptian and Levantine cats were degraded and yielded no useful DNA. Sadly, the DNA extraction from Cyprus cat sample (Shillourokambos) was unsuccessful.
Based on data, sites from Anatolia and Levant had no Clade C, but again we do not have a data to show that IV-C haplotypes were in Egyptian cats before the Classic Era
– Claudio Ottoni
The number of modern cat samples is disappointing, knowing they are probably easier to get compared to the ancient ones.
Lack of historical continuity in many samples is obvious: data from a couple of historical periods is presented but then we have to guess what is there in between.
Researchers believe that IV-C lineage is from Egypt. But does the data support this conclusion?
To get a better idea of the C lineage and Egyptian relation, let’s take a look at samples of Egyptian cats (Figure 4).
Figure 4. Egyptian cat samples used in a study (Supplementary Data 1,2).
All the samples of Egyptian cats came from the same historical period (around 800 BC – 700 CE). We have no idea what genetics Egyptian cats had in earlier historical periods. The large and important chunk of history is lost. While 3 mummies and cats from Oxyrhynchus had haplotype IV-C, one mummy, Berenike and Shenhur cats were IV-A, typical Anatolian. By combining these samples, we can see that both IV-C and IV-A were dominant in Egyptian cats.
These samples tell absolutely nothing whether IV-C is Egyptian or not. In fact, the presence of IV-A haplotype indicates that there was an influx of cats from Anatolia and Levant to Egypt. This can be correlated with studies on human genetics. It is known that Neolithic farmers migrated to Africa and left a genetic imprint on African populations (26). Analysis of genomes of 90 human mummies from Egypt (dated 1388 BC – 426 CE) has also revealed that ancient Egyptians were most closely related to Bronze Age and Neolithic Levantine, Anatolian and European populations (42). Immigrants from these areas had a strong influence on ancient Egyptian genetics. It is entirely possible that the influx of cats from Anatolia and Levant arrived in Egypt with them.
Before the final paper was published Dr. Ottoni was unsure about the lineage C being Egyptian “We do not know whether Clade C existed within Anatolia and Levant before Classic times. Based on data, sites from Anatolia and Levant had no Clade C, but again we do not have a data to show that IV-C haplotypes were in Egyptian cats before the Classic Era (25).”
And yet, the final paper explicitly states that C lineage was Egyptian. As it follows there were “two purported domestication centers.”
Anatolian lineages in Egyptian cats were glossed over with one insignificant sentence: “We also found IV-A* in cat remains from the Roman–Egyptian port of Berenike on the Red Sea and in one Egyptian mummy (Fig. 1a–c), which may hint at an introduction of cats from SWA to Egypt.”
Anatolian lineages in Egyptian cats were glossed over with one insignificant sentence: “We also found IV-A* in cat remains from the Roman–Egyptian port of Berenike on the Red Sea and in one Egyptian mummy (Fig. 1a–c), which may hint at an introduction of cats from SWA to Egypt.”
Did Egyptian cats take over Anatolia and Europe in Roman times?
Anatolian cat lying next to Turkish newspaper with a headline that reads, “Cats from Turkey trace their origins back to Egypt”.
“The fact is that based solely on the mtDNA we cannot tell whether they [Egyptian cats] mixed, we’d need genomic analyses.” Dr. Ottoni said in his email (25).
In an interview with Science magazine, he repeated “It’s still unclear, however, whether the Egyptian domestic cat descends from cats imported from the Near East or whether a separate, second domestication took place in Egypt. Further research will have to show (4).”
Near Eastern cats might have been introduced to Egypt (maybe not even once?), but for this hypothesis to be plausible we need further research or better, extensive genomic analyses. When it comes to the origin of C lineage and second domestication event in Egypt, a belief seems to be sufficient.
Luckily, the data they have from Egyptian and other cats nicely clusters with the historical period which is associated with well know stories and legends that came from writings of Herodotus and Diodorus. A good example is the “Egyptian cat ban” (27, 28):
There’s no reason to believe an independent domestication in Egypt
– Carlos A. Driscoll
“We show that, despite a local ban on cat trading being imposed in Egypt as early as 1700 BC, cats carrying IV-C mtDNA spread to most of the Old World.”
Strangely, they do not consider that C lineage could spread from somewhere else, other than Egypt.
Coincidence or not, but when C lineage became widespread, at the same time the lineage E (at lesser extent D) and species of Asian wildcat Felis silvestris ornata appeared both in Anatolia and Egypt. The occurrence of ornata wildcats in Anatolia may have something to do with the migrations from East, maybe from the areas close to the present day Iran (39). Lineages C, D, E could also be related to these migratory events. We can speculate that Southeastern Anatolia and the Levant could be the homeland of lybica wildcats possessing C lineage. Cats from these areas were represented poorly in this study, there were no samples from southeastern Anatolia which is an interesting area if we consider that the earliest evidence of farming, domestication of animals and crops, was documented there (29).
Where lineages E or D came from is not possible to say.
It is hard to ignore the Anatolian lineages that were found as far as Tanzania, Congo, and Senegal. Authors attempt to explain that the occurrence of lineage A is due to “long-distance maritime routes.” This once again confirms that cats from Anatolia and Levant migrated to the African continent, but little can be said about the chronology of these migrations.
The hypothesis that Egyptian wildcats contributed to the domestic cat genetics is likely but so does the hypothesis that the lineage attributed to Egypt did not come from Egypt. If we put the question about the Egyptian cat origin aside, there is also a possibility that so-called “the second wave” was not from Egypt, or maybe there were no “waves” at all.
We are aware that the authors of this study are not historians, but they do use a literature that features some familiar stories about Egyptian cat origins. They probably tried to match their data with the stories and speculations from the books they cited in a reference list.
There’s no reason to believe an independent domestication in Egypt, said a geneticist Carlos A. Driscoll (4), whose research data laid the foundation for this study. He is right. The study is unable to prove that Egyptians domesticated cats independently yet the interpretations presented in a paper create an impression that it happened with near certainty.
Grumpy Anatolians and friendly Egyptians?
Not so grumpy after all? Anatolian cat from Antakya, Hatay. Photo credit: Sergio Gaitan
Ottoni and his team imply that IV-C haplotypes are of Egyptian origin. This is why they conclude that there were two events of domestication. They say that both Anatolian and Egyptian lineages impacted the genetics of the domestic cat. But as we argued before, they do not provide the sufficient evidence that this is the case. As we will see soon, they offer a quite bizarre explanation of why Egyptian cats were imported to Anatolia and Europe. Since cats have existed in Anatolia and Europe from a long time, we can assume these populations were not small, a significant number of cats from Egypt was necessary to alter the gene pool of cats in such vast areas.
“The Egyptian cat must have been very popular […] the Egyptian cat had properties that made it attractive to humans, presumably acquired during the tightening of the human-cat relationship that developed during the Middle and New Kingdoms and became even stronger afterward. […] it is tempting to speculate that the success of the Egyptian cat is underlain by changes in its sociability and tameness.”
Let’s take a closer look.
They say, IV- C haplotypes became common in Anatolian and European cats since 800 BC. Correct, this agrees with the data given.
Then it is claimed, that these haplotypes are from Egyptian cats. They are unable to prove this with the limited set of data. But let’s say, the lineage is from Egypt. Why would places, where humans and cats lived together for many thousands of years before the Egyptian civilization came into existence, would have a need to import cats from Egypt?
Researchers think there is an explanation for this: Egyptian cats were different. They were more sociable, docile and friendlier towards humans. Therefore, Egyptian cats had an advantage over the “wild” and untamed (?) Anatolian cats. For this reason, cats from Egypt were imported to Anatolia and Europe. As a result, Egyptian cats’ haplotypes/mitotypes IV-C outnumbered the native Anatolian haplotypes/mitotypes IV-A.
There is no way we can access the “tameness” level of Egyptian and Anatolian cats. The iconographic evidence is nice to have, but lack of it is not an evidence for anything. Even though Anatolians did not feature cats in their art, it does not mean that cats from Anatolia were not socialized or that they did not have a close relationship with humans.
Ottoni and his team cite Montague’s study (30) about the discovery of a “domestication gene” in cats (more specifically a region in the genome where such genes could possibly be located). Montague compared cat breeds with a few wildcats (none of which were F. lybica!). This is not a good comparison, knowing that cat breeders control the reproduction of their cats and select the most docile ones. Nothing of this sort of artificial selection could have happened in ancient Egypt, where cats were free to choose their own mates (unless they were caged, but so far nobody proposed this as a possibility).
There is a little doubt that behavioral variation is present in many animal species. The study on rats (31) and experiment on breeding foxes (32) for tameness have shown that individual animals with the proposition to tameness can be selected from the wild population and these individuals are not that rare. We can expect that some wildcats in Anatolia possessed “domestication genes”, making them good candidates for taming.
Yet, socialization may matter more than genetics. Living in farming communities, villages and towns offered a stable supply of food for a wildcat, so wildcats had to be in a close contact with humans. In order to survive cats had to adapt to the new environment by becoming tame. This process probably occurred many times throughout 10,000 years of cat and human history, and still continues in our times.
Are we Egyptomaniacs?
When comes to cats, Anatolia does not have the archaeological richness of that found in Egypt. Mummies from Anatolia are rare, there are no known artifacts with cats, certainly nothing like that Egyptian wall painting of the cat eating a fish, authors chose to include in their paper next to blotched tabby figures (33). By including this image of Egyptian cat as a mackerel tabby representative, researchers emphasized the importance of Egypt in their paper (Figure 5).
Figure 5. Egyptian wall painting of a cat eating a fish under a chair.
We should always be careful when interpreting the art and avoid to romanticize the history. We forget that cats were a part of religious “death cult” in Egypt which sometimes led to cruel practices. It is known that ancient Egyptians broke the necks of many young cats and made them into the mummies (34). This, by today’s standards, would be called an “animal cruelty”. The actual tendency to keep cats as pets in Egypt may have emerged much later (38).
Additionally, it does not seem that other cultures outside Egypt had similar attitudes towards cats – the Bastet cult could be of the local importance. If other cultures did not share similar beliefs regarding cats, it is unlikely they valued them in the same way as Egyptians did. That would be another reason, why outsiders were not interested in taking the Egyptian cats back to their countries.
“Scientists more or less forgot about Egypt after Cyprus find,” said Wim Van Neer, one of the leading researchers (4). Was Egypt really forgotten? Not even close. In fact, most of us have preconceptions about Egyptian involvement in a cat domestication – we automatically assume that Egypt must be a part of the cat’s history in some way or another. This expectation persists in many people minds, including scientists and archaeologists. A study also tries to hold on this idea and justify it with historical speculations, iconography and weak conclusions delivered from an incomplete data set. Sure, it is plausible that researcher’s speculations could be confirmed by the future research, but at the moment when there is not enough support for their conclusions, so the biases favoring Egyptian hypothesis are easily recognizable.
Perhaps, due to its unpopular history, Anatolia had never captivated the imagination of Western Europeans in the same way as Egypt did. Western European fascination with “everything Egyptian” is well documented. This phenomenon is called Egyptomania(34).
Egyptomania has been in existence since Roman times and continues to thrive in a popular culture (movies, music, architecture etc.) today (Figure 6). Mythology and adventurous, romanticized stories about ancient Egypt are very appealing. When you add the mysticism of mummification of cats and the Bastet cult, no wonder cat’s history and its domestication become inseparable from ancient Egyptian civilization in our minds.
Figure 6. Examples of Egyptomania from popular culture: music video by Katy Perry, Hollywood movies Cleopatra (1963) and The Egyptian (1954) include domestic cats or cat symbolism.
Vikings are an insignificant detail in a cat history
We mentioned at the beginning of this article that Vikings are also associated with cats. What role Vikings play in a cat history? Actually, Vikings are an insignificant detail in this study. We may talk about cats accompanying trips of Phoenicians, Greeks, and Romans, but these civilizations do not make such good stories. Just like with Egypt, Vikings and their myths are frequently depicted in a popular culture from TV series to computer games.
The admiration of Vikings could be explained by our ignorance – there is a difference who Vikings really were and who we want them to be (35). We are allowed to recreate our own version of Viking “history.” Scandinavian warrior relationship with cats sounds exciting, and no surprise researchers and media are keen to include Vikings into the cat domestication topic even though it is not a very relevant detail.
Did Ottomans breed blotched tabby cats?
When did coat colors and patterns, other than mackerel tabby, emerge? Researchers were curious about this but only looked for the blotched tabby mutation in their samples. They found this tabby pattern was present only in samples from the Middle Ages, and first occurred in South Western Anatolia which was a part of the Ottoman Empire at that time (Figure 7). Some media sources interpreted this as an example of “selective breeding” (11) for “fancy coats” (13) and “the origin of modern cat shows” (8). It sounds stretched too far: the selective breeding as we know it, is a bit more than a century old, and its beginning is marked by the cat show organized in London in 1871 (40). It’s hard to imagine Ottomans breeding cats for any purpose; and why would they?
Figure 7. Mackerel tabby (grey squares) and blotched tabby (black squares) alleles detected in samples of cats.
Why did researchers choose to look for the blotched tabby? That’s a good question. The study by Kaelin et al. gave a clue to the researchers that the origin of the blotched tabby mutation could be recent (36). Kaelin relied on Linnaeus’ observations that blotched tabbies were frequently depicted on artifacts from the Middle Ages, and this type of art “increased to a sufficiently high frequency […] as characteristic for the domestic cat.” Linnaeus was a renowned taxonomist who gave the blotched tabby a taxonomic name Felis catus– today used in a broader sense to describe all domestic cats (37). So researchers admitted they based their choice of this particular trait because of Kaelin et al., and the latter was inspired by Linnaeus.
Ottoni and his team could also study other color phenotypes. But probably they would fail to detect many of them in a small number of samples they had, and if they did it’s not clear how they would choose to interpret them.
Yet, the researchers still drew conclusions about blotched tabby’s origin. It is understandable. Just like with a “second wave” from Egypt, blotched tabby was a part of their discoveries which they wanted to show off in their publication, nevertheless, it is still problematic. Even in modern cat populations to detect all color variations would need a fairly large sample from different locations, because some color phenotypes may be missing in certain populations.
So did blotched tabbies first occurred in the Middle Ages? We do not know. Dating color phenotypes is difficult. The blotched tabby mutation could possibly be much more ancient than claimed by this study.
Conclusion
This study offers some valuable insights into the domestic cat history. It confirms that cats were first tamed in Anatolia when humans began to practice agriculture. However, widely publicized “second wave” of cats supposedly coming from Egypt cannot be supported by the current evidence.
Where exactly were lybica wildcats tamed first? Where did cats worship by ancient Egyptians come from? How did populations of domestic cat ancestors interact and how they changed throughout different historical periods? These and many other questions still remain unanswered.
While the important fragments of the domestic cat history have already been found, there is a lot of more waiting to be researched and discovered.
Main reference: Ottoni, C., Van Neer, W., De Cupere, B., Daligault, J., Guimaraes, S., Peters, J., … & Bălăşescu, A. (2017). The palaeogenetics of cat dispersal in the ancient world. Nature Ecology & Evolution,1(7), s41559-017.
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