Cat Sense - Part 1
Library

Part 1

CAT SENSE.

How the New Feline Science.

Can Make You a Better.

Friend to Your Pet.

John Bradshaw.

To Splodge.

(19882004).

A Real Cat.

Dogs look up to us: cats look down on us.

-WINSTON CHURCHILL.

When a man loves cats, I am his friend and comrade, without further introduction.

-MARK TWAIN.

Splodge.

Preface.

What is a cat? Cats have intrigued people ever since they first came to live among us. Irish legend has it that "a cat's eyes are windows enabling us to see into another world"-but what a mysterious world that is! Most pet owners would agree that dogs tend to be open and honest, revealing their intentions to anyone who will pay them attention. Cats, on the other hand, are elusive: we accept them on their terms, but they in turn never quite reveal what those terms might be. Winston Churchill, who referred to his cat Jock as his "special a.s.sistant," famously once observed of Russian politics, "It is a riddle, wrapped in a mystery, inside an enigma; but perhaps there is a key"; he might as well have been talking about cats.

Is there a key? I'm convinced that there is, and that it can be found in science. I've shared my home with quite a few cats-and have become aware that "ownership" is not the appropriate term for this relationship. I've witnessed the birth of several litters of kittens, and nursed my elderly cats through their heartbreaking final declines into senility and ill health. I've helped with the rescue and relocation of feral cats, animals that literally wanted to bite the hand that fed them. Still, I don't feel that, on its own, my personal involvement with cats has taught me very much about what they are really like. Instead, the work of scientists-field biologists, archaeologists, developmental biologists, animal psychologists, molecular biologists, and anthrozoologists such as myself-has provided me with the pieces that, once a.s.sembled, begin to reveal the cat's true nature. We are still missing some pieces, but the definitive picture is emerging. This is an opportune moment to take stock of what we know, what is still to be discovered, and, most important, how we can use our knowledge to improve cats' daily lives.

Getting an idea of what cats are thinking does not detract from the pleasures of "owning" them. One theory holds that we can enjoy our pets' company only through pretending that they are "little people"-that we keep animals merely to project our own thoughts and needs onto them, secure in the knowledge that they can't tell us how far off the mark we are. Taking this viewpoint to its logical conclusion, forcing us to concede that our cats neither understand nor care what we say to them, we might suddenly find that we no longer love them. I do not subscribe to this idea. The human mind is perfectly capable of simultaneously holding two apparently incompatible views about animals, without one canceling the other out. The idea that animals are in some ways like and in others quite unlike humans lies behind the humor of countless cartoons and greetings cards; these simply would not be funny if the two concepts negated each other. In fact, quite the opposite: the more I learn about cats, both through my own studies and through other research, the more I appreciate being able to share my life with them.

Cats have fascinated me since I was a child. We had no cats at home when I was growing up, nor did any of our neighbors. The only cats I knew lived on the farm down the lane, and they weren't pets, they were mousers. My brother and I would occasionally catch intriguing glimpses of one of them running from barn to outhouse, but they were busy animals and not over-friendly to people, especially small boys. Once, the farmer showed us a nest of kittens among the hay bales, but he made no special effort to tame them: they were simply his insurance against vermin. At that age, I thought that cats were just another farm animal, like the chickens that pecked around the yard or the cows that were driven back to the barn every evening for milking.

The first pet cat I ever got to know was the polar opposite of these farm cats, a neurotic Burmese by the name of Kelly. Kelly belonged to a friend of my mother's who had bouts of illness, and no neighbor to feed her cat while she was hospitalized. Kelly boarded with us; he could not be let out in case he tried to run back home, he yowled incessantly, he would eat only boiled cod, and he was evidently used to receiving the undivided attention of his besotted owner. While he was with us, he spent most of his time hiding behind the couch, but within a few seconds of the telephone ringing, he would emerge, make sure that my mother's attention was occupied by the person on the other end of the line, and then sink his long Burmese canines deep into her calf. Regular callers became accustomed to the idea that twenty seconds in, the conversation would be interrupted by a scream and then a muttered curse. Understandably, none of us became particularly fond of Kelly, and we were always relieved when it was time for him to head back home.

Not until I had pets of my own did I begin to appreciate the pleasures of living with a normal cat-that is to say, a cat that purrs when it is stroked and greets people by rubbing around their legs. These qualities were probably also appreciated by the first people to give houseroom to cats thousands of years ago; such displays of affection are also the hallmark of tamed individuals of the African wildcat, the domestic cat's indirect ancestor. The emphasis placed on these qualities has gradually increased over the centuries. While most of today's cat owners value them for their affection above all else, for most of their history, domestic cats have had to earn their keep as controllers of mice and rats.

As my experience with domestic cats grew, so did my appreciation of their utilitarian origins. Splodge, the fluffy black-and-white kitten we bought for our daughter as compensation for having to relocate, quickly grew into a large, s.h.a.ggy, and rather bad-tempered hunter. Unlike many cats, he was fearless in the face of a rat, even an adult. He soon learned that depositing a rat carca.s.s on our kitchen floor for us to find when we came downstairs for breakfast was not appreciated, and after that he kept his predatory activities private-without, I suspect, giving the rats themselves any respite.

However brave he was against a rat, Splodge usually kept away from other cats. Every now and then, we would hear the cat-flap clatter as he arrived home in a tearing hurry, and a quick glance out the window would usually reveal one of the older cats in the neighborhood, glaring in the general direction of our back door. He had a favorite hunting area in the park nearby, but kept himself inconspicuous when traveling there and back. His diffidence toward other cats, especially males, was not just typical of many cats; it also exemplified a weakness in social skills that is perhaps the greatest difference between cats and dogs. Most dogs find it easy to get along with other dogs; cats generally find other cats a challenge. Yet many of today's owners expect their cats to accept other cats without question-either when they themselves wish to get a second cat, or when they decide to move, depositing their unsuspecting cat into what another cat thinks is its territory.

For cats, a stable social environment is not enough; they rely on their owners to provide a stable physical environment as well. Cats are fundamentally territorial animals that put down powerful roots in their surroundings. For some, their owner's home is all the territory they need. Lucy, another of my cats, showed no interest in hunting, despite being Splodge's great-niece; she barely strayed more than a dozen yards from the house-except when she came into season and disappeared over the garden wall for hours on end. Libby, Lucy's daughter and born in my home, was as brave a hunter as Splodge had been, but preferred to call the tomcats to her rather than go to them. Even though they were all related and all lived in the same house most of their lives, Splodge, Lucy, and Libby all had distinctive personalities, and if I learned one thing from observing them it was that no cat is completely typical: cats have personalities, just as humans do. This observation inspired me to study how such differences come about.

The transformation of the cat from resident exterminator to companion cohabiter is both recent and rapid, and-especially from the cat's perspective-evidently incomplete. Today's owners demand a different set of qualities from their cats than would have been the norm even a century ago. In some ways, cats are struggling with their newfound popularity. Most owners would prefer that their cats did not kill cute little birds and mice, and those people who are more interested in wildlife than in pets are becoming increasingly vocal in their opposition to the cat's predatory urges. Indeed, cats now probably face more hostility than at any time in the past two centuries. Can cats possibly shake off their legacy as humankind's vermin exterminator of choice, and in just a few generations?

Cats themselves are oblivious to the controversy caused by their predatory natures, but all too aware of the difficulties they encounter in their dealings with other cats. Their independence, the quality that makes cats the ideal low-maintenance pet, probably stems from their solitary origins, but it has left them poorly equipped to cope with many owners' a.s.sumptions that they should be as adaptable as dogs. Can cats become more flexible in their social needs, so that they are unfazed by the proximity of other cats, without compromising their unique appeal?

One of my reasons for writing this book is to project what the typical cat might be like fifty years from now. I want people to continue to enjoy the company of a delightful animal, but I'm not sure that the cat, as a species, is heading in the right direction. The more I've studied cats, from the wildest feral to the most cosseted Siamese, the more I've become convinced that we can no longer afford to take cats for granted: a more considered approach to cat keeping and cat breeding is necessary if we are to ensure their future.

Introduction.

The domestic cat is the most popular pet in the world today. Across the globe, domestic cats outnumber "man's best friend," the dog, by as many as three to one.1 As more of us have come to live in cities-environments for which dogs are not ideally suited-cats have, for many, become the lifestyle pet of choice. About one-third of US households have one or more cats, and they are found in more than a quarter of UK families. Even in Australia, where the domestic cat is routinely demonized as a heartless killer of innocent endangered marsupials, about a fifth of households own cats. All over the world, images of cats are used to advertise all kinds of consumer goods, from perfume to furniture to confectionery. The cartoon cat "h.e.l.lo Kitty" has appeared on more than 50,000 different branded products in more than sixty countries, netting her creators billions of dollars in royalties. Even though a significant minority of people-perhaps as many as one person in five-don't like cats, the majority who do show no sign of relinquishing even a fraction of their affection for their favorite animal.

Cats somehow manage to be simultaneously affectionate and self-reliant. Compared to dogs, cats are low-maintenance pets. They do not need training. They groom themselves. They can be left alone all day without pining for their owners as many dogs do, but they will nonetheless greet us affectionately when we get home (well-most will). Their mealtimes have been transformed by today's pet-food industry from a ch.o.r.e into a picnic. They remain un.o.btrusive most of the time, yet seem delighted to receive our affection. In a word, they are convenient.

Yet despite their apparently effortless transformation into urban sophisticates, however, cats still have three out of four feet firmly planted in their wild origins. The dog's mind has been radically altered from that of its ancestor, the gray wolf; cats, on the other hand, still think like wild hunters. Within a couple of generations, cats can revert back to the independent way of life that was the exclusive preserve of their predecessors some ten thousand years ago. Even today, many millions of cats worldwide are not pets but feral scavengers and hunters, living alongside people but inherently distrustful of them. Due to the astonishing flexibility with which kittens learn the difference between friend and foe, cats can move between these dramatically different lifestyles within a generation, and the offspring of a feral mother and feral father can become indistinguishable from a cat descended from generations of pets. A pet that is abandoned by its owner and cannot find another may turn to scavenging; a generation or two on, and its descendants will be indistinguishable from the thousands of feral cats that live shadowy existences in our cities.

As cats become more popular and ever more numerous, those who revile them continue to raise their voices, but with more venom now than for several centuries before. Cats have never shared the "unclean" tag foisted on the dog and the pig, but despite cats' superficially universal acceptance, a minority of people across all cultures finds cats disagreeable, and as many as one in twenty say that they find them repulsive.2 When asked, few Westerners will admit that they don't like dogs: those who do usually turn out either not to like animals in general or can trace their aversion to a specific experience, perhaps being bitten in childhood.3 Cat-phobia is more deeply seated and less widespread than common phobias of snakes and spiders-phobias that have a logical basis in helping the sufferer to avoid poisonous varieties-but is just as powerful an experience for those who suffer from it.4 Cat-phobics were likely at the forefront of the religious persecution that led to the killing of millions of cats in medieval Europe, and cat-phobia was likely just as common then as it is today. Thus, there can be no guarantee that the cat's popularity will last. Indeed, without our intervention, the twentieth century may turn out to have been the cat's golden age.

Today, spurred on by confessed cat-haters, the cat is coming under attack on the specific grounds that it is a wanton and unnecessary killer of "innocent" wildlife. These voices are most loudly raised in the Antipodes, but are becoming increasingly strident in the UK and the United States. The anti-cat lobby, at its most extreme, demands that cats no longer be allowed to hunt, that pet cats be kept indoors, and that feral cats should be exterminated. Owners of outdoor cats are vilified for supporting an animal that is portrayed as laying waste to the wildlife around their homes. Veterinarians who seek to manage the welfare of feral cats by neutering and vaccinating them, and then returning them to their original territory, have come under attack from within their own profession, with some colleagues charging that this const.i.tutes (illegal) abandonment that benefits neither the cat nor the adjacent wildlife.5 Both sides in this debate admit that cats are "natural" hunters, but cannot agree on how this behavior might be managed. In parts of Australia and New Zealand, cats are defined as "alien" predators introduced from the Northern hemisphere, and are banned from some areas and subject to curfews or compulsory microchipping in others. Even in places where cats have lived alongside native wildlife for hundreds of years, such as in the United States and the UK, their increasing popularity as pets has prompted a vocal minority to press for similar restrictions. Cat owners point to a lack of scientific evidence that pet cats contribute significantly to a population decline of any wild bird or mammal, which are caused instead mainly by the recent proliferation of other pressures on wildlife, such as loss of habitat. Consequently, any restrictions imposed on pet cats are unlikely to result in a resurgence of the species that they supposedly threaten.

Cats themselves are of course unaware that we no longer value their hunting prowess. Insofar as they are concerned, the greatest threat to their subjective well-being comes not from people, but instead from other cats. In the same way that cats are not born to love people-this is something they have to learn when they are kittens-they do not automatically love other cats; indeed, their default position is to be suspicious, even fearful, of every cat they meet. Unlike the highly sociable wolves that were forebears to modern dogs, the ancestors of cats were both solitary and territorial. As cats began their a.s.sociation with humankind some 10,000 years ago, their tolerance for one another must have been forced to improve so that they could live at the higher densities that man's provision of food for them-at first accidental, then deliberate-allowed.

Cats have yet to evolve the optimistic enthusiasm for contact with their own kind that characterizes dogs. As a result, many cats spend their lives trying to avoid contact with one another. All the while, their owners inadvertently compel them to live with cats they have no reason to trust-whether the neighbors' cats, or the second cat obtained by the owner to "keep him company." As their popularity increases, so inevitably does the number of cats that each cat is forced into contact with, thereby increasing the tensions that each experiences. Finding it ever harder to avoid social conflict, many cats find it nearly impossible to relax; the stress they experience affects their behavior and even their health.

The well-being of many pet cats falls short of what it should be-perhaps because their welfare does not grab headlines in the way that dogs' welfare does, or perhaps because they tend to suffer in silence. In 2011, a UK veterinary charity estimated that the average pet cat's physical and social environment scored only 64 percent, with households that owned more than one cat scoring even lower. Owners' understanding of cat behavior scored little better, at 66 percent.6 Without a doubt, if cat owners understood more about what makes their cats tick, many cats could live much happier lives.

Faced with such pressures, cats need not our immediate emotional reactions-irrespective of whether we find them endearing or not-but instead a better understanding of what they want from us. Dogs are expressive; their wagging tails and bouncy greetings tell us in no uncertain terms when they are happy, and they do not hesitate to let us know when they are distressed. Cats, on the other hand, are undemonstrative; they keep feelings to themselves and rarely tell us what they need, beyond asking for food when they're hungry. Even purring, long a.s.sumed to be an unequivocal sign of contentment, is now known to have more complex significance. Dogs certainly benefit from the knowledge of their true natures that can come only from science, but for cats this comprehension is essential, for they rarely communicate their problems to us until these have become too much to bear. Most of all, cats require our a.s.sistance when, as happens far too often, their social lives run awry.

Cats desperately need the kind of research from which dogs have benefited, but unfortunately feline science has not seen the explosion of activity that has recently occurred in canine science. Cats have simply not grabbed the attention of scientists as dogs have. However, the past two decades have provided significant advances, profoundly affecting scientists' interpretations of how cats view the world, and what makes them "tick." These new insights form the core of this book, giving us the first indications on how to help cats adjust to the many demands we now put on them.

Cats have adapted to live alongside people while retaining much of their wild behavior. Apart from the minority that belong to a breed, cats are not humankind's creation in the sense that dogs are; rather, they have coevolved with us, molding themselves into two niches that we have unintentionally provided for them. The first role for cats in human society was that of pest controller: some 10,000 years ago, wild cats moved in to exploit the concentrations of rodents provided by our first granaries, and adapted themselves to hunting there in preference to the surrounding countryside. Realizing how beneficial this was-cats, after all, had no interest in eating grain and plant foods themselves-people must have begun to encourage cats to stay by making available their occasional surpluses of animal products, such as milk and offal. The cats' second role, which undoubtedly followed hard on the heels of the first but whose origins are lost in antiquity, is that of companion. The first good evidence that we have for pet cats comes from Egypt some 4,000 years ago, but women and children in particular may have adopted kittens as pets long before this.

As far as humans are concerned, these dual roles of pest controller and companion have ceased to go hand-in-hand. Although we treasured cats until recently for their prowess as hunters, few owners today express delight when their cat deposits a dead mouse on their kitchen floor.

Cats carry the legacy of their primal pasts, and much of their behavior still reflects their wild instincts. To understand why a cat behaves as it does, we must understand where it came from and the influences that have molded it into what it is today. Therefore, the first three chapters of this book chart the cat's evolution from wild, solitary hunter to high-rise apartment-dweller. Unlike dogs, only a small minority of cats has ever been intentionally bred by people-and furthermore, when there has been deliberate breeding, it has been exclusively for appearance. No one has bred cats to guard houses, to herd livestock, or to accompany or a.s.sist hunters. Instead, cats have evolved to fill a niche brought about by the development of agriculture, from its beginnings in the harvest and storage of wild grains, to today's mechanized agribusiness.

Of course, when the cat first infiltrated our settlements many thousands of years ago, its other qualities did not go unnoticed. Its appealing features, its childlike face and eyes, the softness of its fur, and, crucially, its ability to learn how to become affectionate toward us, led to its adoption as a pet. Subsequently, humankind's pa.s.sion for symbolism and mysticism elevated the cat to iconic status. Popular att.i.tudes toward cats have been profoundly influenced by such connotations: extreme religious views toward cats have affected not only how they were treated, but their very biology-both how they behave and how they look.

Cats have changed to live alongside humans, but we have very different ways of gathering information about and thereby interpreting the physical world that we share. Chapters 4 through 6 examine those differences: humans and cats are both mammals, but our senses and brains work in different ways. Cat owners often underappreciate these differences: we have a natural tendency to a.s.sume that animals perceive the world around us in the same way as we do. Moreover, even in today's world of rationality and science, we still treat the world as if it were sentient, attributing intention to the weather, the earth, and the movements of the stars in the sky. How easy, therefore, it is to fall into the trap of thinking that because cats are communicative and affectionate, they must be, more or less, little furry humans.

Science, however, reveals that cats are anything but. Beginning with the way that every kitten constructs its own version of the world, with consequences that will last its entire lifetime, this part of the book describes how the cat gathers information about its surroundings, especially the way it uses its hypersensitive sense of smell; how its brain interprets and uses that information; and how its emotions guide its responses to opportunities and challenges alike. In scientific circles, it has only recently become acceptable to talk about animal emotions, and one school of thought still maintains that emotions are a byproduct of consciousness, meaning that no animals except humans and possibly a few primates can possibly possess them. However, common sense dictates that if an animal that shares our basic brain structure and hormone systems looks frightened, it must be experiencing something very like fear-probably not in quite the same way that we experience it, but fear nonetheless.

Most (but not everything) of what biology has revealed about the cat's world fits the idea that cats have evolved as predators first and foremost. Cats are social animals too; otherwise, they could never have become pets as well as hunters. The demands of domestication-first of all, the need to cohabitate with other cats in human settlements, and then the benefits of forming affectionate bonds with people-have extended cats' social repertoires out of all recognition compared to those of their wild ancestors. Chapters 7 through 9 explore these social connections in detail: how cats conceive of and interact with other cats and with people, and why two cats may react very differently in the same situation. In other words, we will examine the science of cat "personality."

The book concludes with an examination of the cat's current place in the world, and how this might evolve in the coming decades. Cats are under pressure from many different interests, some well-meaning and others antagonistic. Pedigree cats are still in a minority, and those who breed them are in a position to avoid the practices that have so adversely affected the welfare of pedigree dogs over the past few decades.7 However, the growing fashion for hybrids between domestic cats and other, wild, species of cat, resulting in "breeds" such as the Bengal, can have unintended consequences. We must also ask whether the cat is being inadvertently and subtly altered by those who hold cat welfare closest to their hearts. Paradoxically, the drive to neuter as many cats as possible, with its laudable aim of reducing the suffering of unwanted kittens, may be gradually eliminating the characteristics of the very cats best suited to living in harmony with humankind: many of the cats that avoid neutering are those that are most suspicious of people and the best at hunting. The friendliest, most docile cats are nowadays neutered before leaving any descendants, while the wildest, meanest ferals are likely to escape the attention of cat rescuers and breed at will, thus pushing the cat's evolution away from, rather than toward, better integration with human society.

We are in danger of demanding more from our cats than they can deliver. We expect that an animal that has been our pest controller of choice for thousands of years should now give up that lifestyle because we have begun to find its consequences distasteful or unacceptable. We also expect that we should be free to choose our cat's companions and neighbors without regard for their origins as solitary, territorial animals. Somehow, we presume that because dogs can be flexible in their choice of canine companions, cats will be equally tolerant of whatever relationships we expect them to develop, purely for our convenience.

Until about twenty or thirty years ago, cats kept pace with human demands, but they are now struggling to adapt to our expectations, especially that they should no longer hunt, and no longer desire to roam away from home. In contrast to almost every other domestic animal, whose breeding has been strictly controlled for many generations past, the cat's transition from wild to domestic has-with the exception of pedigree cats-been driven by natural selection. Cats essentially evolved to fit opportunities that we provided. We allowed them to find their own mates, and those kittens that were best suited to living alongside humans, in whatever capacity was required of them at the time, were the most likely to thrive and produce the next generation.

Evolution is not going to produce a cat that has no urge to hunt and that is as socially tolerant as a dog-at least not within a timescale that will be acceptable to the cat's detractors. Ten thousand years of natural selection has provided the cat with enough flexibility to fend for itself when, from time to time, its compact with man breaks down, but not enough to cope with a demand that has grown from nowhere in just a few years. Even for such a prolific breeder as the cat, natural selection would take many generations to move even a token step in this direction. Only deliberate, carefully considered breeding can produce cats that are well suited to the demands of tomorrow's owners, and that will be more acceptable to cat haters.

We can do much to improve the lot of cats today. Better socialization of kittens, better understanding of what environments cats really need, more deliberate intervention in teaching cats to cope with situations that they find distressing-all these can help cats to adjust to the demands we now make of them, and can also deepen the bond between cat and owner.

Cats are in many ways the ideal pet for the twenty-first century, but will they be able to adapt to the twenty-second? If they are to continue to remain in our affections-and the persecution they have received in the past indicates that this is hardly a given-then some consensus must emerge among cat welfare charities, conservationists, and cat fanciers on how to produce a type of cat that checks all the boxes. These changes must be guided by science. Initially, the way forward will be for cat owners and the general public alike to understand better where cats came from and why they behave as they do. At the same time, owners can rehabilitate the cat's fraying reputation by learning how to channel their cat's behavior, not only to discourage them from hunting, but also to make them happier in themselves. In the longer term, the emerging science of behavioral genetics-the mechanics of how behavior and "personality" are inherited-will allow us to breed cats that can better adapt to an ever-more crowded world.

As history shows, cats can fend for themselves in many ways. However, they cannot face what society now demands of them without human a.s.sistance. Our understanding of cats must start with a healthy respect for their essential natures.

CHAPTER 1.

The Cat at the Threshold Pet cats are now a global phenomenon, but how they transformed themselves from wild to domestic is still a mystery. Most of the animals around us were domesticated for prosaic, practical reasons. Cows, sheep, and goats provide meat, milk, and hides. Pigs provide meat; chickens, meat and eggs. Dogs, our second favorite pet, continue to provide humans with many benefits beside companionship: help with hunting, herding, guarding, tracking, and trailing, to name but a few. Cats are not nearly as useful as any of these; even their traditional reputation as rodent controllers may be somewhat exaggerated, even though, historically, this was their obvious function as far as humankind was concerned. Therefore, in contrast to the dog, we have no easy answers as to how the cat has insinuated itself so effectively into human culture. Our search for explanations starts some ten millennia ago, when cats probably first arrived at our doorsteps.

Conventional accounts of the domestication of the cat, based on archaeological and historical records, propose that they first lived in human homes in Egypt about 3,500 years ago. This theory, however, has recently been challenged by new evidence coming from the field of molecular biology. Examination of differences between the DNA of today's domestic and wild cats has dated their origins much earlier, anywhere between ten thousand and fifteen thousand years ago (8,000 and 13,000 years BCE). We can safely discount the earliest date in this range-anything earlier than about 15,000 years ago makes little sense in terms of the evolution of our own species, since it is unlikely that stone-age hunter-gatherers would have had the need or resources to keep cats. The minimum estimate, 10,000 years, presumes that domestic cats are derived from several wild ancestors that came from several different locations in the Middle East. In other words, the domestication of the cat happened in several widely separated places, either roughly contemporaneously or over a longer period of time. Even if we a.s.sume that cats started to become domesticated around 8,000 years BCE, this leaves us with a 6,500-year interval before the first historical records of domestic cats appear in Egypt. So far, few scientists of any kind have studied this first-and longest-phase in the partnership between human and cat.

The archaeological record for this period, such as it is, is not very illuminating. Cats' teeth and fragments of bones dating between 7,000 and 6,000 BCE have been excavated around the Palestinian city of Jericho and elsewhere in the Fertile Crescent, the "cradle of civilization" that extends from Iraq through Jordan and Syria to the eastern sh.o.r.es of the Mediterranean and Egypt. However, these fragments are uncommon; moreover, they could well have come from wild cats, perhaps killed for their pelts. Rock paintings and statuettes of catlike animals from the following millennia, discovered in what is now Israel and Jordan, could conceivably depict domesticated cats; however, these cats are not depicted in domestic settings, so they may well be representations of wild cats, possibly even big cats. Yet even if we a.s.sume that these pieces of evidence all refer to early forms of the domestic cat, their very rarity still must be explained. By 8,000 BCE, humankind's relationship with the domestic dog had already progressed to the extent that dogs were routinely buried alongside their masters in several parts of Asia, Europe, and North America, whereas burials of cats first became common in Egypt around 1,000 BCE.1 If cats were indeed domesticated pets during this time, we should have far more tangible evidence of that relationship than has been uncovered.

Our best clues on how the partnership between man and cat began come not from the Fertile Crescent, but instead from Cyprus. Cyprus is one of the few Mediterranean islands that have never been joined to the mainland, even when that sea was at its lowest level. Consequently, its animal population has had to migrate there by flying or swimming-that is, until humans started to travel there in primitive boats some 12,000 years ago. At that point, the Eastern Mediterranean had no domesticated animals, with the likely exception of some early dogs, so the animals that made the crossing with those first human settlers must have been either individually tamed wild animals or inadvertent hitchhikers. Therefore, while we cannot possibly tell whether ancient remains of cats on the mainland are from wild, tame, or domesticated animals, cats could clearly have reached Cyprus only by being deliberately transported there by humans-a.s.suming, as we safely may, that cats of that era were as averse to swimming in the ocean as today's cats are. Any remains of cats found there must be those of semidomesticated or at least captive animals, or their descendants.

On Cyprus, the earliest remains of cats coincide with and are found within the first permanent human settlements, some 7,500 years BCE, making it highly likely that they were deliberately transported there. Cats are too large and conspicuous to have been accidentally transported across the Mediterranean in the small boats of the time: we know very little about seagoing boats from that period, but they were likely too small to conceal a stowaway cat. Moreover, we have no evidence for cats living away from human habitations on Cyprus for another 3,000 years. The most likely scenario, then, is that the earliest settlers of Cyprus brought with them wildcats that they had captured and tamed on the mainland. It is implausible that they were the only people to have thought of taming wildcats, so capturing cats and taming them were likely already an established practice in the Eastern Mediterranean. Confirming this, we also have evidence for prehistoric importations of tamed cats to other large Mediterranean islands, such as Crete, Sardinia, and Majorca.

The most likely reason for taming wildcats is also evident from the first settlements on Cyprus. Right from the outset, these habitations, like their counterparts of the time on the mainland, became infested by house mice. Presumably these unwanted mice were stowaways, accidentally transported across the Mediterranean in sacks of food or seed corn. The most likely scenario, therefore, is that as soon as mice became established on Cyprus, the colonizers imported tame or semidomesticated cats to keep them under control. This might have been ten years or a hundred years after the first settlements were established-the archaeological record cannot reveal such small differences. If this is correct, it suggests that the practice of taming cats to control mice was already entrenched on the mainland as long as 10,000 years ago. No firm evidence for this is ever likely to be found, because the ubiquitous presence of wildcats there makes it impossible to tell whether the remains of a cat, even if found within a settlement, are those of a truly wild cat that had died or been killed when hunting there, or of a cat that had lived there for most or all of its life.

Whatever its exact origins, the tradition of taming wildcats to control vermin continued into modern times in parts of Africa where domestic cats are scarce and wildcats easy to obtain. While traveling the White Nile in 1869, the German botanist-explorer Georg Schweinfurth found that his boxes of botanical specimens were being invaded by rodents during the night. He recalled: One of the commonest animals hereabouts was the wild cat of the steppes. Although the natives do not breed them as domestic animals, yet they catch them separately when they are quite young and find no difficulty in reconciling them to a life about their huts and enclosures, where they grow up and wage their natural warfare against the rats. I procured several of these cats, which, after they had been kept tied up for several days, seemed to lose a considerable measure of their ferocity and to adapt themselves to an indoor existence so as to approach in many ways to the habits of the common cat. By night I attached them to my parcels, which were otherwise in jeopardy, and by this means I could go to bed without further fear of any depredations from the rats.2 Like Schweinfurth, those much earlier explorers who first brought wildcats to Cyprus would almost certainly have found that they had to keep the cats tethered. If allowed to run free, the cats would have quickly escaped and wreaked havoc on the native fauna, which up to that point would have had no experience with a predator as formidable as a cat. We know that this is what eventually happened. Several centuries after human settlement, cats indistinguishable from wildcats spread throughout Cyprus and remained there for several thousand years.3 Most likely, only the cats that were confined to the grain stores would have stayed there to help the early settlers to rid those stores of pests: The others would have left to exploit the local wildlife. The descendants of these escapees may have been captured and even eaten from time to time, since broken cat bones have been found at several other Neolithic sites on Cyprus, as well as those of other predators such as foxes and even domestic dogs.

The practice of taming wildcats to control vermin was probably prompted by the emergence of a new pest in the early granaries, the house mouse (Mus musculus); indeed, the histories of these two animals are inextricably interwoven. The house mouse is one of more than thirty species of mouse found worldwide, but the only one that has adapted to living alongside humans and exploiting our food.

House mice have their origins in a wild species from somewhere in northern India that was in existence possibly as long as a million years ago, certainly well before the evolution of humankind. From there they spread both east and west, feeding on wild grains, until some reached the Fertile Crescent, where they eventually encountered the earliest stores of harvested grain: mouse teeth have been found among stored grain dating back 11,000 years in Israel, and a 9,500-year-old stone pendant carved in the shape of a mouse's head was found in Syria. Thus began an a.s.sociation with humankind that continues to the present day. Humans not only provided an abundance of food that mice could exploit, but our buildings also provided both warm, dry places to build nests and protection from predators such as wildcats. Mice that could adapt to these living conditions thrived, while those that could not died out: today's house mice rarely breed successfully away from human habitation, especially where there are wild compet.i.tors, such as wood mice.

Humans also provided house mice with a way to colonize new areas. Mice from the southeastern part of the Fertile Crescent, what is now Syria and northern Iraq, were accidentally transported, presumably in grain being traded between communities, throughout the Near East, up to the eastern sh.o.r.es of the Mediterranean, and then to nearby islands such as Cyprus.

The first culture to be bedeviled by house mice was that of the Natufians, who by extension are the people most likely to have initiated the cat's long journey into our homes. The Natufians inhabited the area that now comprises Israel-Palestine, Jordan, southwestern Syria, and southern Lebanon from about 11,000 to 8,000 BCE. Widely regarded as the inventors of agriculture, they were initially hunter-gatherers like other inhabitants of the region; soon, however, they began to specialize in harvesting the wild cereals that grew abundantly all around them, in a region that was significantly more productive then than it is today. To do this, the Natufians invented the sickle. Sickle blades found at Natufian settlements still show the glossy surfaces that could have been produced only from scything through the abrasive stems of wild grains-wheat, barley, and rye.

The early Natufians lived in small villages; their houses were partly belowground, partly above, with walls and floors of stone and brushwood roofs. Until about 10,800 BCE, they rarely planted cereals deliberately, but over the next 1,300 years a rapid change in climate, known as the Younger Dryas, brought about a significant intensification in field-clearing, planting, and cultivation. As the amounts of harvested grain increased, so did the need for storage. The Natufians and their successors probably used storage pits built out of mudbrick and constructed like miniature versions of their houses. It was probably this invention that triggered the self-domestication of the house mouse, which, moving into this rich and novel environment, thereby became humankind's first mammalian pest species.

As the numbers of mice grew, they must have attracted the attentions of their natural predators, including foxes, jackals, birds of prey, the Natufians' domestic dogs, and, of course, wildcats. Wildcats had two advantages that set them apart from other predators of mice: they were both agile and nocturnal, well adapted to hunting in the near-darkness when the mice became active. However, if these wildcats had been as frightened of man as their modern counterparts are, it is difficult to imagine how they would have exploited this new rich source of food. Almost certainly, therefore, the wildcats in the region inhabited by the Natufians were less wary than those of today.

We have no evidence that the Natufians deliberately domesticated the cat. Like the mouse before it, the cat simply arrived to exploit a new resource that had been created by the beginnings of agriculture. As Natufian agriculture became more complex, involving both an increasing array of crops and the domestication of animals such as sheep and goats, and as agriculture extended to other regions and cultures, so the opportunities available for cats multiplied. These were not pet cats as we know them today; rather, the cats that exploited these concentrations of mice would have been more like today's urban foxes-capable of adapting to a human environment, but still retaining their essential wildness. Domestication was to come much later.

We know surprisingly little about the wildcats of the Fertile Crescent and surrounding areas (see box on page 8, "The Evolution of the Cats"). The archaeological record indicates that 10,000 years ago several species lived in the region, all of which would have been attracted by concentrations of mice. We know that later on, the ancient Egyptians kept tame jungle cats, Felis chaus, in considerable numbers; jungle cats, though, are substantially heavier than wildcats, weighing between ten and twenty pounds, and large enough to kill young gazelle and chital. Although their normal diet includes rodents, they may have been too obtrusive to get regular access to granaries. Alternatively, they may simply have been temperamentally unsuited to living alongside man. We do have evidence that the Egyptians tried to tame and even train them as rodent controllers, but apparently without any lasting success.

The Evolution of the Cats Every member of the cat family, from the n.o.ble lion to the tiny black-footed cat, can trace its ancestry back to a medium-sized catlike animal, Pseudaelurus, that roamed the steppes of central Asia some 11 million years ago. Pseudaelurus eventually went extinct, but not before unusually low sea levels had allowed it to migrate across what is now the Red Sea into Africa, where it evolved into several medium-sized cats, including those we know today as the caracal and the serval. Other Pseudaelurus traveled east across the Bering land bridge into North America, where they eventually evolved into the bobcat, lynx, and puma. Some 2 to 3 million years ago, following the formation of the Panama isthmus, the first cats crossed into South America; here they evolved in isolation, forming several species not found anywhere else, including the ocelot and Geoffroy's cat. The big cats-lions, tigers, jaguars, and leopards-evolved in Asia and then spread into both Europe and North America, their present-day distributions but a tiny relict of where they used to roam a few million years ago. Remarkably, the distant ancestors of today's domestic cats seem to have evolved in North America about 8 million years ago, and then migrated back into Asia some 2 million years later. About 3 million years ago, these began to evolve into the species we know today, including the wildcat, the sand cat, and the jungle cat; a separate Asian lineage, including Pallas's cat and the fishing cat, also began to diverge at about this time.4 Jungle cat Coeval with them were sand cats, Felis margarita, large-eared nocturnal animals that hunt by night, using their acute hearing. They are, moreover, comparatively unafraid of humans, and hence might be thought good candidates for taming and domestication. However, they are made for life in deserts-the pads on their feet are covered in thick fur to protect them from the hot sand-so few would have found themselves near the first stores of grains: the Natufians generally built their villages in wooded areas.

Sand cat As civilization spread eastward through Asia, so it would have come into contact with other cat species. At Chanhudaro, a town built by the Harappan civilization close to the Indus River in what is now Pakistan, archaeologists found a 5,000-year-old mudbrick imprinted with a cat's foot, overlapped by that of a dog. As the newly made brick was drying in the sun, the cat appears to have run across it, closely followed by a dog, possibly in hot pursuit. The footprint is larger than that of a domestic cat, and its webbed feet and extended claws identify it as a fishing cat, Felis viverrina, found today from the Indus basin eastward and south to Sumatra in Indonesia (though not in the Fertile Crescent). As its name implies, the fishing cat is a strong swimmer and specializes in catching fish and aquatic birds. Although it will also take small rodents, it is difficult to see how it would switch to a diet consisting predominantly of mice, so it too is an unlikely candidate for domestication.

Manul Farther afield, we know of at least two other species of cat that came in out of the wild to prey on the vermin that plagued humankind's food stores. In Central Asia and ancient China, the local wildcat, the manul (or Pallas's cat, named after the German naturalist who first categorized it) was occasionally even tamed and deliberately kept as a rodent controller. The manul has the s.h.a.ggiest coat of any member of the cat family, so long that its hair almost completely obscures its ears. In pre-Columbian Central America, meanwhile, an otter-like cat, the jaguarundi, was probably also kept as a semi-tame pest controller. None of these species have ever become fully domesticated, neither are any of them included in the direct ancestry of today's house cats.

Jaguarundi Out of all these various wild cats, only one was successfully domesticated. This honor goes to the Arabian wildcat Felis silvestris lybica, as confirmed by their DNA.5 In the past, both scientists and cat-fanciers have suggested that certain breeds within the domestic cat family are hybrids with other species-for example, the Persian's fluffy feet are superficially similar to the sand cat's, and its fine coat is somewhat like that of a manul. However, the DNA of all domestic cats-random-bred, Siamese, or Persian-shows no trace of these other species, or indeed any other admixture. Somehow, the Arabian wildcat alone was able to inveigle itself into human society, outcompeting all its rivals, and eventually spreading throughout the world. Although the qualities that gave it this edge are not easy to pin down, they probably occurred in combination only in the wildcats of the Middle East.

The wildcat Felis silvestris is currently found throughout Europe, Africa, and central Asia, as well as western Asia, the area where it probably first evolved. Like many predators, such as the wolf, it is now found only in isolated and generally remote areas where it can avoid persecution from man. This has not always been the case. Five thousand years ago, wildcats were evidently regarded as delicacies in some areas; the rubbish pits left by the "lake dwellers" of Germany and Switzerland contain many wildcat bones.6 The cats must have been abundant at the time; otherwise, they could hardly have been trapped in such large numbers. Over the centuries they became less common, displaced by the felling of their forest habitat for agriculture, and forced farther into the woods by development and loss of habitat. The invention of firearms led to wildcats being hunted to extinction in many areas. During the nineteenth century, various European countries, including the UK, Germany, and Switzerland, cla.s.sified them as vermin, due to the harm they supposedly caused both wildlife and livestock.7 Only recently, due to the establishment of wildlife reserves and a more informed att.i.tude to the important role that predators play in stabilizing ecosystems, are wildcats returning to areas such as Bavaria, where they have not been seen for hundreds of years.

The wildcat is now divided into four subspecies or races. These are the European forest cat Felis silvestris silvestris, the Arabian wildcat Felis silvestris lybica, the Southern African wildcat Felis silvestris cafra, and the Indian desert cat Felis silvestris ornata.8 All these cats are rather similar in appearance, and all are capable of interbreeding where their ranges overlap. A possible fifth subspecies is the very rare Chinese desert cat Felis bieti, which according to its DNA split off from the main wildcat lineage about a quarter of a million years ago. It's possible that these cats actually form a separate species, as no hybrids are known to exist, but they live in such a small and inaccessible region-part of the Chinese province of Sichuan-that this may be due to lack of opportunity rather than physical impossibility.

Wildcats from different parts of the world differ markedly in how easily they can be tamed. Domestication, moreover, can start only with animals that are already tame enough to raise their young in the proximity of people. Those offspring that are best suited to the company of humans and human environments are, perhaps unsurprisingly, more likely to stay and breed there than those that are not; the latter will most likely revert to the wild. Over several generations, this repeated "natural" selection will, even on its own, gradually change the genetic makeup of these animals so that they become better adapted to life alongside people. It is also likely that, at the same time, humans will intensify that selection, by feeding the more docile animals and driving away those p.r.o.ne to bite and scratch. This process cannot start without some genetic basis for tameness existing beforehand, and in the case of wildcats, this is far from evenly distributed. Today, some parts of the world have little raw material for domestication, while others seem more promising.

The distribution of the subspecies of wildcat We know, for example, that the four subspecies of wildcat differ in how easy they are to tame. The European forest cat is larger and thicker-set than a typical domestic cat, and has a characteristic short tail with a blunt, black tip. This aside, it looks from a distance much like a domestic striped tabby-a distant glimpse is all that most people are likely ever to get, however, for it is among the wildest of animals. This is largely due to its genetics, and not the way it is raised: those few people who have tried to produce tame forest cats have met with precious little but rejection. In 1936, natural and wildlife photographer Frances Pitt wrote: It has long been stated that the European wildcat is untameable. There was a time when I did not believe this. . . . My optimism was daunted when I made acquaintance with Beelzebina, Princess of Devils. She came from the Highlands of Scotland, a half-grown kitten that spat and scratched in fiercest resentment. Her pale green eyes glared savage hatred at human-beings, and all attempts to establish friendly relations with her failed. She grew less afraid, but as her timidity departed, her savagery increased.9 Pitt then went on to obtain an even younger male kitten, in the hope that Beelzebina had been too old to be socialized when first found. That she named this new kitten Satan perhaps suggests how difficult he was to handle from the outset. As he grew stronger and more confident, he became impossible to touch; he would take food from the hand, but would spit and growl while doing so, and then quickly back away. However, he was not pathologically aggressive-he just hated people. While he was still young, Pitt introduced him to a female domestic kitten, Beauty, toward whom he was "all gentleness and devotion." When she was let out of the cage in which he had to be kept, "this distressed him sorely. He rent the air with harsh cries, for his voice, though loud, was not lovely." Beauty and Satan produced several litters of kittens, all of which had the characteristic appearance of forest cats. Some, despite being handled from an early age, grew to be as savage as their father; others were more sociable toward Pitt and her parents, though all remained very wary of unfamiliar people. Pitt's experiences with Scottish wildcats seem to be typical: Mike Tomkies, the "Wilderness Man," was also unable to socialize his two hand-raised wildcat sisters, Cleo and Patra, which he kept at his remote cottage on the sh.o.r.es of a Scottish loch.10 We know little about the Indian desert cat, but it is reputedly difficult to tame. This subspecies is found to the south and east of the Caspian Sea, southward through Pakistan and into the northwestern Indian states of Gujarat, Rajasthan, and the Punjab, and eastward through Kazakhstan into Mongolia. Its coat is usually paler than that of the other wildcats, and is blotchy rather than tabby in pattern. Like other wildcats, it will occasionally base itself near farms, attracted by the concentration of rodents, but it has never taken the next step to domestication-acceptance of humans. We have records from Harappa of tamed caracals, a medium-sized, long-limbed cat with characteristic tufted ears, and jungle cats, to add to the fishing cat that left its footprint there; but we find no indications of any Indian desert cats. For a long time biologists and cat fanciers alike thought that Siamese cats could be a mixture of domestic and Indian desert cat, the progeny of interbreeding between early domestic cats and local wildcats somewhere around the Indus valley. However, scientists have not found the characteristic DNA signature of the Indian desert cat in any examples of the Siamese and related breeds, which instead are ultimately derived from the wildcats of the Middle East or Egypt-there are no silvestris wildcats in Southeast Asia, so the original Siamese cats must have arrived from the west as fully domesticated animals.

The wildcats of South Africa and Namibia-"caffre cats"-are likewise genetically distinctive. They migrated south from the original wildcat population in northern Africa about 175,000 years ago, around the same time as the ancestors of the Indian desert cat migrated east. It is unclear where the boundary between the Southern African and Arabian wildcats lies-no wildcat's DNA has yet been characterized from any part of Africa except Namibia and the Republic of South Africa. The wildcats of Nigeria are shy, aggressive, and difficult to tame; those of Uganda are sometimes more tolerant of people, but many do not look like typical wildcats-which in that area have distinctive red-brown backs to their ears-and are probably hybrids, their domesticated genes accounting for their friendly behavior. Most of the street cats in the same region show signs of some wildcat in their ancestry, so the distinction between wildcat, street cat, and random-bred pet is blurred in many parts of Africa.

The wildcats of Zimbabwe-presumably belonging to the Southern African subspecies-are a case in point. In the 1960s, naturalist and museum director Reay Smithers kept two hand-reared female wildcats, Goro and Komani, at his home in what was then Southern Rhodesia.11 Both were tame enough to be let out of their pens, though only one at a time, since they would fight whenever they met. Once, Komani disappeared for four months, finally reappearing one evening in the beam of Smithers's flashlight: "I called my wife, to whom she is particularly attached, and we sat down while she softly called the cat's name. It must have taken a quarter of an hour before Komani suddenly responded and came to her. The reunion was most moving, Komani going into transports of purring and rubbing herself against my wife's legs."

Such behavior is identical to that of a pet cat being reunited with its owner, and the similarities with pet cats did not end there. Both Goro and Komani were affectionate toward Smithers's dogs, rubbing themselves on their legs and curling up in front of the fire with them. Every day they demonstrated their affection for Smithers himself by an effusive display of typical pet cat behavior.

These cats never do anything by halves; for instance when returning from their day out they are inclined to become super-affectionate. When this happens, one might as well give up what one is doing, for they will walk all over the paper you are writing on, rubbing themselves against your face or hands; or they will jump on your shoulder and insinuate themselves between your face and the book you are reading, roll on it, purring and stretching themselves, sometimes falling off in their enthusiasm and, in general, demanding your undivided attention.

This may be the behavior of a typical hand-reared "caffre cat," but it is more likely that Goro and Komani, while undoubtedly wildcats in terms of their markings and their hunting ability, nevertheless contained some DNA from interbreeding with pet cats somewhere in their ancestry. The extent of hybridization between wildcats and domestic cats in South Africa and Namibia was recently revealed by DNA sequences from twenty-four supposed wildcats, eight of which bore the telltale signs of partial descent from domestic cats. In a survey of zoos in the United States, the UK, and the Republic of South Africa, I found that ten out of twelve South African wildcats displayed affectionate behavior toward their keepers, and of these, two would regularly rub and lick them.12 This kind of behavior strongly suggests that the latter were hybrids, while those that could not be handled at all were probably genuine wildcats. The eight that were moderately affectionate might have been either.

Hybridization between wildcats and domestic cats is not confined to Africa. In one study, five out of seven wildcats collected in Mongolia carried traces of domestic cat DNA; only two were "pure" Indian desert cat. In my survey of cats in zoos, I found that out of a dozen cats of this subspecies kept in captivity, only three had ever spontaneously approached their keepers, and only one had ever rubbed on its keeper's leg. From the proportions found in the DNA results, it seems highly likely that all of these were hybrids, even though they all looked like typical Indian desert cats. In the same study of wildcat DNA, almost a third of apparent "wildcats" sampled in France had some ancestry from domestic pets.13 With the advent of DNA technology, it is easy to detect hybridization when the local wildcats are genetically distinct from domestic cats-as they are in southern Africa, central Asia, and western Europe alike. Defining what is wild and what is a hybrid is much more problematic in places where domestic cats and wild cats are genetically almost identical, as they are around the Fertile Crescent, home of the Arabian wildcat.

The Arabian wildcat lybica is not only the most similar to domestic cats, it is also probably the nearest living representative of the first Felis silvestris, all the other subspecies having evolved hundreds of thousands of years ago-a consequence of small numbers of animals migrating east, south, or west from the species' origin in the Middle East. The wildcats in Africa north of the Sahara are also probably lybica, but their DNA has not yet been tested to confirm this. Like all wildcats, the Arabian/North African wildcat has a "mackerel" striped tabby coat, varying in color from gray to brown-darkest in forest-dwelling animals, palest in those that live on the edges of deserts. It is generally larger and leaner than a typical domestic cat, and both its tail and legs are especially long; indeed, the front legs are so long that when it sits, its posture is characteristically upright, as depicted by the Ancient Egyptians in statues of the cat G.o.ddess Bast. While generally nocturnal and therefore rarely seen, it is not particularly rare. Although it is widely claimed that the Arabian wildcat's kittens, if hand-raised, become affectionate toward people, most eyewitness accounts come from central or southern Africa, and therefore probably refer to cafra rather than lybica. The explorer Georg Schweinfurth procured his tame wildcats in what is now Southern Sudan, roughly where the ranges of lybica and cafra merge together, and the most northerly location in Africa to yield reliable accounts of tamable wildcats.

Very little is known of the behavior of genuine lybica wildcats, either in the Middle East or northeast Africa. In the 1990s, conservationist David Macdonald radio-collared six wildcats on the Thumamah reserve in central Saudi Arabia. All except one kept their distance from human activity: the sixth, however, "often wandered into the vicinity of the pigeon house [in Thumamah town] and would often be found sleeping with the domestic cats in the yard of one of the houses. On one occasion he was seen mating with a [domestic] cat."14 Apart from showing just how easily hybridization between wild and domestic cats can occur, these and other observations shed little light on whether the wildcats in this part of the world might have been easy to tame, thousands of years ago.

The Arabian wildcat-Felis lybica Traci