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  Cats can live as cherished domestic pets or as tolerated wild or half-wild animals living in proximity to human dwellings. Domestic pets generally rely on their human benefactors for food and shelter, while the wild/half-wild cats live as self-sufficient hunters or scavengers.

  Biologists consider cats to be "exploitive captives" - captives of humans, but exploiting this relationship to their own ends rather than suffering from it. Throughout its history, cats have also been symbiotes where both humans and cats gain from the relationship - humans gain rat-catchers while cats gain shelter and supplementary feeding. Elsewhere, where rodent control is not an issue, they are commensals - present in the same place as man and dining with him, probably consuming leftovers and providing a form of garbage collection (the food scraps may otherwise have attracted vermin or disease). Because of these roles, cats and humans have a long history together. Archaeological evidence for this relationship dates to around 8000 years ago.

  In his contribution to Frances Simpson's "The Book of the Cat" (1903), H C Brooke ("Some Foreign Cats") opted not to go into detail on wild felines, but briefly mentioned the Egyptian Cat (Felis maniculata) and the European Wild cat (F catus) (note: these are not the Latin names by which the cats are known today) and wrote:

  "It might reasonably be imagined that our common cat was derived from the last-named, considering that at one time it was a common animal all over England as well as on the continent. The untameable ferocity of this variety - which is probably the least amenable of all living creatures - has doubtless prevented its ever having been domesticated and the high value which, as we learn from old writings, was a common animal in England, plainly show that F catus was not the ancestor of F domestica, although the two will freely interbreed. Many years ago, the old Spanish wild cat which used to be kept at the Zoological Gardens in the so-called aviaries, now occupied by the civets, mated with his cage mate - a tortoiseshell-and-white queen - and of these cross-bred kittens both Sir Claud Alexander and the writer of these lines [Brooke] possessed specimens. It is usually assumed that the Egyptian or Caffre cat is the progenitor of the majority of the domestic cats […] it is found over the whole of Africa, and it is quite easy to understand how, with its eminently tameable disposition, it gradually spread over Europe. Our so-called Abyssinian cats […] bear a very striking resemblance to this handsome variety of cat. "

  In 1907, the taxonomist, Pocock, described two types of domestic cat F torquata (striped tabby) and F catus (blotched tabby) although F catus was elsewhere used to describe the European wildcat. Linnaeus assigned the name F catus to the domestic blotched tabby. The domestic cat has also been known as F domestica.

  The accepted ancestor of the domestic cat (Pocock 1907) is the Arabian and African form of the wildcat, Felis lybica (F lybica). The European wildcat (including the Scottish wildcat) is F silvestris. These two wildcats interbreed easily so they are now considered related subspecies properly called F silvestris silvestris (a robust form adapted to colder climes) and F silvestris lybica (J Clutton Brock, A Natural History of Domesticated Mammals, 106-112, 1987). The prehistory of domestic cats is complicated and it is suggested that species other than F s lybica were involved e.g F s silvestris (European wildcat) and F ornata (Asiatic wildcat).

  Felis lybica has a very large geographic range and several subspecies whose appearance varies. These subspecies can interbreed to produce offspring whose appearance is intermediate between the parents. Generally speaking, moving from north to south there is a gradation of coat thickness, intensity of ground colour and the amount of "tabby" markings (Robinson 1991), with northern subspecies being more heavily built, thicker furred with a rounded-tipped tail and darker, more distinct tabby markings than southern subspecies. The African wildcat has a lighter build, less distinct markings and thin, tapering tails and varies from rusty, through sandy and tawny to grey with faint tabby markings.

  In 1951, Pocock described 26 subspecies of wildcat though many were probably variants or intermediates since only 4 groups are recognised today. Weigel (1961) and Hemmer (1978) described four groups: the "forest" cats (silvestris) of Europe, the Caucasus and Asia Minor; the "steppe" cats (ornata) of South and Central Asia; the "tawny" cats (lybica) of Africa and the Middle East; and the familiar domestic cat (catus).

  As late as 1996, and in spite of the hybridisation at London zoo which H C Brooke reported (1903), some researchers claimed that F catus was not closely enough related to F s silvestris for them to be able to produce fertile hybrids. In fact these hybridize naturally in Scotland and the local sub-species of F silvestris, F s grampia (Scottish Wildcat) is in danger of being supplanted by hybrids. In the Amur region of Russia there is a domestic breed (the Ussuri) believed to be descended from F catus x F s silvestris wildcats, however this is based on appearance only.

  A species credited with partial ancestry of the domestic cat (especially of those in India) is F chaus, the jungle cat or marsh cat. F chaus was present in Ancient Egypt and is possibly depicted in Ancient Egyptian art. In addition, there is evidence that F chaus was kept alongside F s lybica in Egyptian temples during the time that cats were domesticated. The two species interbreed successfully, but there is, as yet, no compelling evidence for past hybridization between the two species.

  Felis chaus produces fertile offspring with domestic cats, but its contribution to the ancestral gene pool is considered to be minimal. Some biologists believe that the input of F chaus is greater than currently admitted and that this has largely been ignored (R Tabor, Cats - The Rise of the Cat, p16, 1991). Tabor states that F catus x F chaus hybrids are far more docile than F catus x F s silvestris hybrids; suggesting that F chaus contributed the genes which made early domestic cats humanisable. However, F catus x F s lybica hybrids are also more docile than F catus x F s silvestris hybrids so the docilising genes could equally have come from F s lybica. If anything, the hybrid offspring of domestic cats (or African Wild cats) and European Wild cats take after the European Wild cat (i.e. non-docile) and will move away from man.

  Tabor suggests other species which may have contributed to the mix of genes in the domestic and which may account for some of the traits not found in F s lybica, F s silvestris or F chaus. These species include the golden cat (F temminckii) which occurs between south China and Sumatra and which, he believes, may have contributed towards the oriental type cats. It is sometimes suggested that Persian longhairs may be partly descended from Pallas's cat (F manul) found in Iran through to western China, although there is no skeletal evidence for this. Others suggestions include the leopard cat (F bengalensis) from eastern Siberia through to South-east Asia, and the sand cat (F margarita) found from the Sahara to Baluchistan.

  Their general appearance may suggest that they contributed to the evolution of the domestic cat, but detailed studies of skulls, skeletons and anatomical features do not support this. Domestic cats are anatomically and skeletally similar to F s lybica. More recently, chromosome counts show that it is feasible for Pallas's cat, the Leopard cat and the Asian Golden cat to have contributed in the past, but the domestic cat is most similar in type to the African wildcat. It is equally likely that variations in appearance (colour, fur length) are due to genetic mutations.

  It is certain that domestic cats evolved from one or more sub-species of F silvestris. It is possible, but not proven, that other species have been involved such that the domestic cat is a complex hybrid of several species.

  While Brooke correctly identified the Egyptian or Caffre (now known as F s lybica) cat as progenitor of "the majority of the domestic cats", he incorrectly believed that the domestic cats around the world were descended from the smaller wild cats of the countries in question. According to the naturalist Blyth, quoted by Brooke, this resulted in the two varieties of Indian domestic cats: the fulvous (ticked) variety derived from the Jungle cat (F chaus) and the spotted varieties supposedly derived from the Leopard cat, Desert cat and Rusty Spotted cat. To illustrate this belief in different races of domestic cats in India, Brooke wrote of an Indian cat acquired by his wife from a London shoemaker who had acquired from a sailor who had stolen it from a hotel in Bombay.

  "Very curious and handsome is the Indian cat[…] His most striking peculiarities are the length and slenderness of his limbs, the extreme shortness of his coat, and his thin and tapering tail […] His ears are small, but as a kitten they were of enormous size, and with his long and pointed head gave him a most weird experience. The voice of this cat is very variable, and far more resembles the raucous call of the Siamese than the voice of any European cat." In retrospect, it is probably that the cat was an Oriental rather than a variety descended from indigenous wildcats, but in Brooke's time only the only recognised Oriental was the Siamese. The variety had probably reached Bombay the same way it left - on shipboard, probably from Thailand or the Malay Peninsula. The Siamese of that time being far less extreme than the modern variety; the illustration shows the Indian cat as similar to an Abyssinian.

  Why should the African wildcat have proven domesticable while its more northern cousins did not? As well as colouration among subspecies across its range, temperament also varies. The more northerly F silvestris is generally intractable even if hand-reared. F lybica is generally more amenable to human presence. This was obviously a factor in the early domestication of the cat.

  African wildcats hunt a variety of prey animals, but rarely scavenge carrion. Unlike feral domestic cats, which sometimes form colonies, African wildcats are solitary. Domestic cats will often eat carrion and will form colonies where there is a rich food supply e.g. rubbish dumps, farmyards, but prefer to remain solitary where prey is more thinly distributed (Liberg and Sandell 1988). Studies in Saudi Arabia indicate that wildcats continue to remain solitary while feral cats formed colonies around a garbage dump.

  However, African wildcats do have some ability to co-operate since captive female wildcats have been seen to assist mothers in supply food for kittens (Smithers 1983), a behaviour observed in feral cat colonies. Early domestication is believed to be due to humans attracting rodents which attracted cats. Those wildcats which best tolerated the presence of other cats would have been better able to exploit the food supply and would have produced young which inherited their more sociable tendencies. In addition to becoming more sociable, cats also evolved to exploit humans by scavenging carrion and food scraps.

  Remains of cats have been found from the Pre-pottery Neolithic of Jericho (circa 7000 BC) though these may have been hunted animals or the equivalent of modern feral cats, attracted to food scraps and tolerated at best. Tamed cats have possibly lived in association with humans far earlier than archaeological and historical records imply. Cat remains (bones, teeth) are often retrieved from prehistoric sites, but it is impossible to know whether these were companions or prey (pelts, meat).

  Initially, cats were probably tolerated by humans because they killed the mice and rats that were attracted to human food stores. The cats themselves may have been eaten or hunted for their pelts. Later, humans would have deliberately encouraged the presence cats as rat-catchers instead of hunting them.

  In Cyprus, African Wild cat remains dated to around 5000 BC have been found. This is interesting because the cat must have been transported to Cyprus by humans. Cyprus has never had any indigenous wildcats. The bones of cats, mice, and humans were found together and all three of these species first appear on Cyprus at the same time. This suggests that humans brought mice to the island of Cyprus by accident and the cats on purpose to control the mice.

  Nineteenth century explorers of North Africa have recorded that local people caught and reared African Wild Cats as pets, keeping them around their huts and enclosures. Similar events have been reported (right up to modern times) in North African and Arabia.

  The Azande tribe of southern Sudan live in tolerant proximity with the African Wild cat, though they haven't domesticated it. The cats are content to find shelter and scavenge food around the tribal settlements and show little fear of people. In this it is much like the urban fox in Britain - tolerates proximity with humans but not tame.

  The relative docility of the African Wild cat and its ability to leave close to people without being held in captivity, suggest that the cat itself moved in with people exploiting shelter and food supplies in and around human settlements.

  According to J A Baldwin, there seems to be little question that European domesticated cats came out of Egypt between 3,000 BC and 2,000 BC (J A Baldwin, Anthropos 70:428-448, 1975). There are no confirmed records of domesticated cats from the Old Kingdom (2686-2181 BC). The earliest Ancient Egyptian evidence of the presence of cats dates from about 2500 BC and there is evidence of primitive domestication around 2000 BC, however the Egyptians also depict many non-domestic species. There is little pictorial evidence to suggest that cats were fully domesticated until the New Kingdom, 18th Dynasty (circa 1600 BC).

  A small number of F chaus skulls have been recovered from Egyptian burial sites, but no skulls with features intermediate between F s lybica and F chaus have yet been reported. Some Egyptian paintings depict cats with features consistent with F chaus in a marshland setting.

  The role of cats in Ancient Egyptian religion is well documented so there is no need to describe it in detail here. Cats were kept (and bred) in temples where they would have been in close proximity with humans and dependent upon them for food. This would have contributed to their domestication as the tamest cats would have thrived best in such closed and continued proximity with people. Numerous cat mummies have been found; many appear to have been sacrifices and cats would have been reared in large numbers for this purpose. Because of their religious status, as well as their rat-catching abilities, cats were also kept by Egyptian households.

  Paintings would seem to suggest that cats were used in hunting of wildfowl although this is debatable since cats are not generally good retrievers. Later they were depicted in paintings as symbols of fertility and/or domestic harmony.

  Ancient sites have been discovered in the Indus Valley (now Pakistan) dating back to 6000 BC. The Indus had a flourishing civilization with several major cities comprising 40,000 to 50, 000 inhabitant at the time of the Egyptian kingdoms. Domestication of cats may have followed a parallel route to that known to have occurred in Ancient Egypt.

  Indus valley sites (e.g. Harrapa, Mohenja-daro) indicate the presence of granaries and the domestication of animals (Jarrig & Meadow, Scientific American 243:122-133, 1980) such as cats, dogs, goats, sheep and possibly elephants. At its peak, this civilization stretched across the whole of Sindh, Baluchistan, Punjab, Northern Rajasthan, Kathiawar and Gujarat. The cities were far more advanced than their counterparts in prehistoric Egypt, Mesopotamia or elsewhere in Western Asia. In common with most other contemporary civilizations, agriculture was the backbone of the Indus economy and the granaries would have attracted rodents which in turn attracted cats.

  Bones of F catus have been found at some sites in the Indus valley in situations which indicate domestication. There is evidence of cat tracks in mud-and-straw bricks used at these sites (much like cat tracks from where cats walk across drying cement). Whether the F catus bones date back as far as 6000 BC wasn't reported. Studies of Asian sites dating back to 5000 BC shows the domestication of animals in this area well before 3000 BC. Archaeological studies of villages established on the coast of the Bay of Bengal have also shown cat remains associated with domestication.

  Historical and archaeological data and genetic evidence (genetic divergence between populations), suggests an origin for the domestic cats of Pakistan in western Asia that predates the Harrapan civilization. The similarities between the Indus and Ancient Egyptian civilizations make it probable that there was a west-central Asian centre for cat domestication independent of Egypt (M Ahmad, B Blumberg, M. F Chudary, Journal of Heredity 71:323-330, 1980).

  The Indus cities had commercial links with Afghanistan, Persia, Egypt, Mesopotamia and the Samaritans. Cats would probably have travelled between Ancient Egypt and the Indus valley with Phoenician traders sometime around 2000 BC, allowing the two populations to interbreed. After 2000 BC, for unknown reasons, the Mohenjo-daro and Harappan culture slowly declined.

  The first human cities appeared around 5000 BC (Catal Huyuk, Turkey). Civilization began much earlier in warm climes (Egypt, Indus valley, South America, Mediterranean) than in northern parts of Europe which were still recovering from a cold period. This put the tractable species of wildcat into close proximity with man, especially where they hunted rats attracted to city granaries. It is conceivable that cat species were tamed in other places where humans built cities.

  The Mohics of Peru were known to worship the cat as a god of copulation and fertility. It is possible that small species of cat were tamed and kept as pets or in temples. The Jaguarundi is believed to have been kept as a pet although it was tamed rather than domesticated.

  Cheetah have been used for hunting, but they were tamed and trained rather than domesticated. Servals have also been tamed and kept as pets. The larger species (Lions, Leopards, Puma) have been kept as pets, but (barring a few exceptional individuals) have an unreliable temperament in captivity and their large size made them dangerous. A few people have kept semi-tame Scottish Wildcats (F silvestris grampia), but these generally revert to wildness as they mature as do modern F s grampia x F catus hybrids.

  The domestic cat was taken to the Americas by European settlers. Some were ship's ratters, polydactyls being preferred for this task which accounts for the high incidence of polydactyly on the eastern coast of the United States (i.e. the original colonies) and a few eastern coastal locations in Canada. Others were pets which accompanied settlers. It is possible that cats accompanied the original Viking settlers, however no evidence of feline occupation remains.

  There have been reports of early colonists finding domestic cats already in North America and it was, for a while, supposed that the cat may have interbred with native lynx. The size difference between the two species is great enough to prevent natural interbreeding and the greyish creatures with banded tails which were reported by early naturalists were more likely to have been racoons.

  In South America there are two cat species which show a high degree of tameability in captivity: Geoffroy's Cat and the Margay. Both have recently been hybridized with domestic cats; the Margay to produce the "Bristol" (later incorporated into the Bengal breed) and the Geoffroy's Cat to produce the Safari. This has caused some naturalists to wonder why these two species were apparently never domesticated.

  The most likely reason is that they are forest cats and that forest peoples did not form static societies like those in Egypt and Pakistan. They were primarily hunter-gathers and did not build granaries and therefore had no use for ratters to protect their grain. There are reports that Jaguarundi kittens were occasionally tamed as pets, but there was no concerted effort to domesticate the Jaguarundi.

  The small cats in regions where Central and South American peoples did build cities were probably not amenable to domestication. Hence no small cat species was domesticated in the Americas.

  Although Ancient Egyptians forbade the export of cats, by 1700 BC the cat was depicted in domestic poses in Israel. By 1400 BC domestic cats were present in Greece. By 1000 BC cats had travelled northwards across the Mediterranean aboard ships (possibly with Phoenician traders) and from there they spread along trade routes. The cat travelled eastward to China and Japan (where it protected silkworm cocoons from rats). The Romans regarded cats as rare and exotic pets, preferring the mongoose for vermin control. By 500 BC, domestic cats seem to have been familiar in southern Europe.

  The cat may have arrived in England with the Phoenicians who traded for tin in Cornwall, though it is most likely the Romans who first brought cats with them some time before 4 AD. Cat footprints have been discovered in clay tiles at Caesaromagus (Chelmsford, Essex). Further cats arrived with the Vikings.

  From then on, cats travelled alongside man in ships, valued as rodent controllers who protected ships' supplies and as lucky mascots. They travelled to America with colonists; the preference for polydactyl cats as lucky shipboard mascots is reflected in the high numbers of polydactyls in the first settled areas on the eastern US coast. In the US, the cat may possibly hybridize with F rufus (Bobcat) and has been deliberately hybridized with F geoffroyii (Geoffroy's cat), F viverrina (Fishing Cat) and F serval (Serval). Future generations of some varieties of F catus will therefore be even more complex hybrids than their predecessors.

  There is some archaeological evidence that the Vikings briefly settled on the American Atlantic coastline around 1000 AD and it has been suggested that domestic cats accompanied them. Count de Buffon's Natural History (1767) suggests that wild cats existed in pre-colonial America and that a hunter brought a wild cat to Christopher Columbus. The cat was described as brown/grey creature, of normal size for a domestic cat and with a long tail.

  It has been suggested that the cat was either it was a European domestic cat and that these had survived unchanged over the centuries, or that it was a domestic x F rufus hybrid. The colouration is consistent with either a Bobcat (or hybrid) or an Abyssinian-type cat. However, the term "cat" has been used over the centuries to describe cat-like non-felids (e.g. Pine Martens were once known as Tree Cats) so the creature described might well have been a non-felid. The absence of domestic cat bones among the archaeological evidence suggests that the domestic cat did not reach America until the first European settlers took them there.

  The cat penetrated Australia with Europeans in 1788, though aboriginal histories indicate that cats were already present on parts of the Australian coast as shipwreck survivors. There are no indigenous species of feline in Australia.

  The Egyptian genesis is still the prevalent theory, but the recent discovery of evidence of domestication at Asian sites as early as 4000 BC to 5000 BC opens the question of independent domestication of cats among other ancient peoples. China's ancient archaeology is still relatively unknown and there is the possibility that Chinese species of small cat may have been domesticable.

  The fact that cat remains have been found in various ancient cities and settlements in places other than Egypt make it feasible that small cat species have always lived in or near human settlements, probably in a semi-tame state, attracted there by shelter and by food. This made the cat, particularly F s lybica, ripe for domestication. Domestication may have occurred simultaneously in several distinct regions with Egypt being the best known example.

  Whether other species have been involved in the evolution of the domestic cats cannot be determined with current DNA analysis technology.

  Is the modern domestic cat still basically a wild animal?

  The proliferation of feral cat colonies demonstrates that modern domestic cats retain enough of their wild ancestry to successfully revert to the wild state and to create self-sustaining populations of wholly wild cats, identical to their domestic cousins in every way except for temperament and, over a suitable time period, a return to the wild type as human-selected traits are removed through natural selection. They remain genetically similar enough to F s grampia and F s lybica to form fully fertile hybrids with these wild species.

  This does not mean that every individual cat can revert to the wild state. Many pedigree cats have been bred into forms unsuited to the wild state - too short noses which prevent hunting, excessive fur which mats and becomes a liability, too little fur so that hypothermia is a hazard, excessive docility or skeletal deformities. Other individuals have had survival traits surgically removed i.e. declawing. In addition, the "wild state" in many areas is full of man-made hazards.

  Many behaviours of pet cats are manifestations of their wild instincts e.g. scent marking, hunting toys. Many outdoor-going cats display a full range of wild behaviours e.g. hunting and marking/protecting territory. In indoor cats, instincts frustrated by close confinement can be redirected into problem behaviours which conflict with their attraction as a pet e.g. middening, attacking ankles. It is understandable that humans want to keep pet cats out of harm's way, but unless the captive environment is enriched, the cat acts out its natural instincts as best it can. Some cats retain more wild behaviours than others which is why not all cats are suited to the indoor-only life forced upon them by the human environment.

  Whether we like to admit it or not, our domestic cat companions are essentially still wild at heart and in their genes and instincts; something which keeps pet behaviourists in business.