From Wildcat to House Cat: The Domestication of the Cat

Cats are one of the most popular house pets on the planet. They can adapt to small living spaces, such as apartments, and are relatively easy to care for. Cats also rule the internet; they are present in many memes, and cat videos are one of the most prevalent videos on YouTube. The question, however, remains, how did the lovable mini house panther come to be; were they first domesticated in Egypt, or in the Near East, and how were they domesticated; did they migrate into farming areas because of the available rodents to eat, or were they attracted by trash; and did they have preadaptive genes that made them easier to domesticate? These and many more questions can be answered with research.

Introduction - Domestication

What is domestication exactly? According to Charlotte Cameron-Beaumont, author of Evidence Suggesting Preadaptation to Domestication Throughout the Small Felidae “The criteria for complete domestication, permanent isolation from the wild species, and human control of breeding, territory and food supply” (2002). Cats are only semi-domesticated, because some breeding populations are not separated from wildcats; and because humans do not provide food or control their breeding (Montague, M.J. et al. 2014). However, pedigree breeds, such as Siamese and Persian, are considered completely domesticated because they are under complete human control so the breed meets the domestication standards. The criteria for domestication are not met for the large populations of cats that have become feral. These feral cats select their own mates, scavenge for their own food, and compete for territories (Cameron-Beaumont, C. et al. 2002). Because of those traits, they are not considered domesticated. Some researchers say the African wildcat, Felis silvestris lybica, may have been meant for domestication because it is docile towards humans. This docile behavior distinguishes it from other Felis silvestris subspecies. The process of cat domestication was very slow progress (Linseele, V. et al. 2007).

The process of domestication should not be confused with taming, they are two different things where one affects an individual versus a group of animals. Taming is the conditioned behavioral modification of an individual; domestication is the permanent genetic modification of a bred lineage that leads to, among other things, a heritable predisposition toward human association. And domestics do not have to be ‘tame’ in the behavioral sense look at the Spanish fighting bull for example. Undomesticated animals can be tame, look at a hand-raised cheetah for example (Driscoll, C. A., Macdonald, D. W., et al. 2009). Cats are considered very unlikely candidates for domestication. Cats are solitary hunters and will fight other cats of the same sex to defend territories. Cats are obligate carnivores, meaning they can only digest meat, while most domestic animals eat plants. (Driscoll, C. A., Clutton-Brock, J. et al. 2009). Another reason why cats might not be considered fully domesticated is that they do not perform direct tasks like dogs can, their purpose is, even as mousers, debatable (Driscoll, C. A., Macdonald, D. W., et al. 2009). Cats most likely chose to live among humans because of the many opportunities they found (Driscoll, C. A., Clutton-Brock, J. et al. 2009).

All domesticates have one thing in common; they are all very tolerant of people (Driscoll, C. A., Macdonald, D. W., et al. 2009). F. s. lybica is the only subspecies of the wildcat that was domesticated, but why? Evidence suggests that other subspecies of wildcats are less tolerant of people such as the European wildcat and the Chinese mountain cat. (Driscoll, C. A., Clutton-Brock, J. et al. 2009). Most domesticates were behaviorally preadapted to domestication.

After 10,000 B.P., cat domestication arose after humans built houses, farms, and settlements (Driscoll, C. A., Macdonald, D. W., et al. 2009). Felis silvestris catus, the ‘domestic’ cat, is the only member of its Family that was successfully domesticated. The cats' domestication process should have been hindered for a few reasons as mentioned before: it is an obligate carnivore, and they do not form large social groups because of being extremely territorial, in addition, they are extremely agile, and they are wary of man (Cameron-Beaumont, C. et al. 2002). Knowing about the domestication of cats brings the question of where did the domestication of Felis catus first begin? Comment by Veronica Lovelace:

The History of the Cat; Where Cats Were First Thought to be Domesticated

Egypt

Scholars once believed that the ancient Egyptians were the first to keep cats as pets, possibly as early as 3,600 years ago (Driscoll, C. A., Clutton-Brock, J. et al. 2009). The exact place and date of the cat's domestication are still undocumented, but their domestic status seems to have been reached by the 12th dynasty, during the Middle Kingdom period of Egypt, when cats begins to appear often in Egyptian art. However, there is an older painting of a cat with a collar around its neck from a tomb in Saqqarah, dated to the 5th dynasty (Linseele, V. et al. 2007). Some of these tomb paintings show cats positioned under chairs, sometimes eating from bowls, and they are often collared. The majority of these paintings show that cats were a common occurrence in Egyptian households (Driscoll, C. A., Clutton-Brock, J. et al. 2009). Many authors have written about the popularity Egyptian house cats gained by the end of the 1st millennium B.C. Some authors have also mentioned that the consumption of cat meat was considered a crime and that it was illegal to export cats outside of the country (Linseele, V. et al. 2007).

The Egyptians then took the love of cats to a whole new level. Nearly 2,900 years ago, cats began to represent the Goddess Bastet, which was the official deity of Egypt at that time. Millions of house cats were sacrificed, mummified in Bastet’s name, and buried in her sacred city, Bubastis (Driscoll, C. A., Clutton-Brock, J. et al. 2009). Cats were one of the most frequently mummified animals in Egypt. Studies on the cat mummies have shown that Felis silvestris, the Wildcat, was the main felid species mummified. Occasionally, the Jungle cat, Felis chaus, and the Serval, Leptailurus serval were also mummified. Other studies have shown that the Egyptian domestic cat was larger than their extant wild species. It is thought, however, that the large size of the domestic cat mummies is possible because of the special status and good nourishment they were given (Linseele, V. et al. 2007). The excessively large number of cat mummies found in tombs would suggest that the Egyptians were, for the first time in history, breeding domestic cats and not just harvesting wild populations (Driscoll, C. A., Clutton-Brock, J. et al. 2009). About 3,000 years ago, even though the cat trade was prohibited in Egypt, cats arrived in Greece and later arrived in Europe (Hu, Y. et al. 2014). However, the cats seem to have spread around the world faster after Egypt came under Roman rule when cats and religion were dissociated (Linseele, V. et al. 2007).

Cyprus

In 2004 on the Mediterranean island of Cyprus, a discovery of possibly the earliest findings of humans keeping cats as pets were found by Jean-Denis Vigne of the National Museum of Natural History in Paris and his colleagues (Driscoll, C. A., Clutton-Brock, J. et al. 2009).

This archaeological evidence was dated back to 9,500 years ago, and shows that humans coexisted with cats (Montague, M. J. et al. 2014). In a shallow grave with an assortment of items, an adult human was laid to rest, and just 40 centimeters away in its own little grave a young cat’s body was arranged in the same way as humans (Driscoll, C. A., Clutton-Brock, J. et al. 2009). This evidence suggested that the cats were tame when brought to Cyprus. The burial of the cat points to having some sort of bond with the human. The burial of the cat also points to it having a role as an individual as well (Linseele, V. et al. 2007). Cats are not native to most of the Mediterranean islands, therefore people must have brought the cats from the adjacent Levantine coast by boat. The burial of the cat and the human, along with cats being transported to Cyprus shows that there was a special relationship between cats and humans nearly 10,000 years ago beginning in the Middle East (Driscoll, C. A., Clutton-Brock, J. et al. 2009).

Near East - Fertile Crescent

James Henry Breasted invented the term Fertile Crescent by characterizing the region by its cultural and ecological features that were present during its early civilization. The Fertile Crescent is a region that is bound by the Mediterranean sea to the coast of the Levant and includes the Mesopotamian plains and most of the Taurus mountains. The Fertile Crescent does not include Egypt.(Driscoll, C. A., Macdonald, D. W., et al. 2009). About 9.500 years ago, cats entered human society around the time people first started farming in the Middle East (Grimm, D. 2014). For many years, it has been assumed that wildcats moved into these agricultural settlements because of their grain storages attracted rodents which wildcats liked to prey upon. Wildcats also liked to take advantage of the settlement's trash heaps that provided easy year-round food supplies (Hu, Y. et al. 2014). Comment by Veronica Lovelace:

Eventually, Wildcats that were the most tolerant of humans moved into Fertile Crescent villages. The people of these villages saw that the wildcats did little harm and noticed that they were actually beneficial because they disposed of rodents and snakes. Villagers eventually encouraged wildcats to stay within the settlements. Agriculture eventually spread beyond the Fertile Crescent and with it, tame members of F. s. Lybica followed behind (Driscoll, C. A., Clutton-Brock, J. et al. 2009).

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China

Around 2,000 years ago, domestic cats appeared in Chinese settlements. However, a rare discovery of felid remains was found in Shaanxi, China. This finding dated back to 5,300 years ago, and showed that cats were present in early Chinese agricultural villages. Most interestingly, this discovery is outside of the range of F. s. lybica. If wildcats were domesticated in the Fertile Crescent more then 5,000 years ago cats showed up in northern China, then it is predicted that cats were transported from western Asia by intricate trading systems similar to how other animals ended up in parts of eastern Asia (Hu, Y. et al. 2014). Soon, isolated groups of domestic Oriental cats began to change from a process known as genetic drift, which gave these cats distinctive coat colors. These traits created by genetic drift were neither beneficial nor harmful and later became fixed in these populations of Oriental cats. Thus the Siamese, the Birman, the Korat, and many other “natural breeds” were born from this process. Many of these cat breeds were described by Buddhist monks in a book called Tamara Maew (meaning “Cat Poems”) that dated back to 1350 A.D. (Driscoll, C. A., Clutton-Brock, J. et al. 2009).

Comparing the House Cat to the Wildcat

It is not certain if all types of domestic cats descended from one species of wildcat, Felis silvestris, but many think this to be now true. The species Felis silvestris is found in many places in the Old World. Wildcat populations are found in Scotland all the way to South Africa, and as far east as Mongolia. In the year 2000, scientists were finally able to tell which wildcat populations turned into a domestic cat by a gathering of 979 DNA samples. These samples were collected from different wildcat and domestic cat individuals from southern Africa, Azerbaijan, Kazakhstan, Mongolia, and the Middle East. If the samples of DNA from the wildcats populations distinguished them from one another and domestic cat DNA closely matched one of the wildcat populations, then scientists could find where domestication first started (Driscoll, C. A., Clutton-Brock, J. et al. 2009). It was discovered that the wildcat DNA of the subspecies F. s. lybica, collected from the deserts of Israel, Saudi Arabia, and the United Arab Emirates, is identical to domestic cat DNA. This means that the domestic cat was first domesticated in the Fertile Crescent, and it is only grouped with the wildcat F. s. lybica (Driscoll, C. A., Clutton-Brock, J. et al. 2009). Comment by Veronica Lovelace:

However, the question remains, are domesticated cats a separate species from their wild counterparts? According to the Biological Species Concept, the answer is “no”. If given the opportunity, domestic cat populations will breed with wildcat populations and they will produce viable fertile offspring (Driscoll, C. A., Macdonald, D. W., et al. 2009). This hybridization, however, is a serious conservation issue in parts of Europe (O’Connor, T. P. 2007). In many of Scotland’s populations of European wildcats, hybridization has been detected (Witzenberger, K. A. et al. 2014). Furthermore, it is difficult to distinguish a purebred wildcat from a feral domestic cat because of the common mackerel tabby phenotype (Driscoll, C. A., Menotti-Raymond, M., et al. 2007). On top of that, modern data from archaeological samples proves that it is impractical to expect differences in biometric measurements between the two animals as well (O’Connor, T. P. 2007). Unless genetic screening is involved, hybrids are virtually impossible to identify (Witzenberger, K. A. et al. 2014). The average domestic cat has a few morphological differences from wildcats, but they mostly retain the wild body style. These differences are slightly shorter legs, a smaller brain, and as Charles Darwin noted, a longer intestine, which may have been an adaptation to scavenging kitchen scraps (Driscoll, C. A., Clutton-Brock, J. et al. 2009).

Was F. s. lybica Domesticated by Commensal Means?

Current thinking emphasizes wildcat domestication as a mutualistic symbiotic relationship between humans and cats. It is thought that cats followed a commensal pathway, which led to domestication (Hu, Y. et al. 2014). It has been suggested that cat domestication happened in several steps. In the first step, wildcats were more than likely drawn to agricultural villages in the Near East by mice that were feeding on the grain storage. In the second step, wildcats began to pray on the commensal rodents in the villages and then may have become commensal themselves. The last step is that humans may have noticed the presence of the wildcats and realized that they were preying upon the mice that ate their food storage. Humans either tolerated or may have encouraged the wildcats to stay in their villages. Eventually, this relationship may have resulted in domestication (Linseele, V. et al. 2007)(Driscoll, C. A., Clutton-Brock, J. et al. 2009). In northern China, a similar process occurred. Ancient Chinese millet farming, along with millet storage attracted mice. The mice in turn attracted wildcats, which led to cats becoming useful in early Chinese agricultural villages as well (Hu, Y. et al. 2014). Comment by Veronica Lovelace:

Rodents more than likely attracted cats to early farming villages, but the trash piles outside of the villages were even easier pickings, and they provided a year-round food supply for resourceful felines (Driscoll, C. A., Clutton-Brock, J. et al. 2009). The first domestic cats were a product of natural selection because early agricultural settlements would more than likely not have purposely sought out the wildcat to make them a pet (Driscoll, C. A., Macdonald, D. W., et al. 2009).

Was F. s. lybica Domesticated by Genetic Predisposition?

Scientists have recently begun to identify genes that may have helped wildcats to become domesticated (Grimm, D. 2014). The domestic cat's genome was found to contain 19,493 protein-coding genes and 1,855 noncoding RNAs (Montague, M.J. et al. 2014). Researchers were able to uncover 13 genes that changed when cats became domesticated. Based on recent studies on knockout mice, a few of these identified genes may play a role in cognition, including fear response and the ability to learn new behaviors when given food rewards (Grimm, D. 2014). This means that because humans were being associated with food, docility may have unintentionally been selected for, and this may have been a major force that altered the first domestic cat genomes (Montague, M.J. et al. 2014). This high-quality sequencing of the cat genome may eventually reveal how other animals became domesticated (Grimm, D. 2014).

A research team also identified a few other genes in the cat genome that code for glutamate receptors. These receptors are associated with learning and memory. Five other genes in the domestic cat genome were found to influence the migration of neural crest cells, these stem cells affect everything from skull shape to coat color in a developing embryo (Grimm, D. 2014).

It is suggested that changes in these neural crest-related genes may have been the cause of the tameness cats developed during their domestication. These findings may also explain why many other domestic animals share common traits such as certain pigmentation patterns and smaller brain sizes, which was first noted by the famous Charles Darwin (Montague, M.J. et al. 2014) (Grimm, D. 2014).

Composite short tandem repeat genotypes have shown that wildcats from the Near East, along with domestic and feral cats, all fall within a large monophyletic group (Driscoll, C. A., Menotti-Raymond, M. et al. 2007). Studies of today’s wildcat and domestic cat mitochondrial DNA show that the ancient Near Eastern wildcat subspecies, Felis silvestris lybica, are their maternal ancestors (Hu, Y. et al. 2014). On top of this finding, hybridization has caused many domestic cat mtDNA haplotypes to be found within indigenous wildcat haplotypes in many wildcat populations around the world (Driscoll, C. A., Menotti-Raymond, M. et al. 2007). All of these genetic changes in the wildcat's genome helped it to become domesticated.

Conclusion

This research shows that cat domestication has happened over many years because of commensal and preadaptive genes. When the first agricultural villages started to develop in the Fertile Crescent, rodents were attracted to the grain storages and trash heaps that these settlements produced. Felis silvestris lybica was attracted to the rodents and trash heaps in these villages due to them being easily obtainable food sources. Certain individuals of Felis silvestris lybica had preadaptive genes that made them tamer which made them prime subjects for domestication. When the ancient people of these villages saw that the cats were beneficial to them, they encouraged the cats to stay in their villages. Many years later, Felis silvestris lybica turned into the domestic cat, Felis catus, that we know and love today.

Researching the process of domestication can help gather information on evolutionary biology, animal welfare science, and conservation biology. Others are researching whether domestic cats have bred with Asian wildcats, or if they themselves have ever been domesticated at some point in time. Some research is being done on ancient felide archaeological finds because there seems to be a lack of research on them compared to studies on ancient dog archaeological finds.

The Cat Fancier’s Association, along with the International Cat Association acknowledge around 30 to 40 cat breeds. Hybrids of different cat breeds are being recognized by the Associations such as the Bengal cat, which is a hybrid between the domestic cat, Felis catus, and the Leopard cat, Prionailurus bengalensis. Other hybrids are also being developed such as the Chausie breed, a hybridization between the Felis chaus and Felis catus, and the Savannah breed, a hybridization between Felis catus and Leptailurus serval. Most of these cat breeds are not being developed for an actual purpose like dogs have been bred for, but rather for aesthetic reasons. This could lead to animal welfare concerns, as many pet owners may not know how to properly take care of these majestic animals.

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