Cat Sense Part 7

A reduced motivation to hunt and kill prey is just one of several factors that will enable cats to adapt better to twenty-first-century living. Allowing a little anthropomorphism: if cats could write themselves a wish list for self-improvement, a set of goals to allow them to adapt to the demands we place on them, it might look something like this: *To get along better with other cats, so that social encounters are no longer a source of anxiety.

*To understand human behavior better, so that encounters with unfamiliar people no longer feel like a threat.

*To overcome the compulsion to hunt even on a full stomach.

The corresponding requests from owners: *I'd like to have more than one cat at a time, and for my cats to be company-not just for me, but also for one another.

*I wish my cat didn't disappear into the bedroom to urinate on the carpet every time I have visitors.

*I wish my cat didn't bring gory "presents" through the cat-flap.

We currently know of two ways to achieve these goals. First, we can train individual cats to change how they interpret and react to their surroundings. The advantage of this approach is that its effects would be immediate; the disadvantage is that it must be repeated for each successive generation of cats. Second, because the cat's genome is not yet fully domesticated, there is still scope to genetically adapt their behavior and personalities to twenty-first-century lifestyles. The benefits of selective breeding toward our goals will become apparent only after several decades, but these changes would be permanent.

Cats are intelligent, and (up to a point) adaptable animals, so we can achieve some of these goals through directing cats' learning-providing them with the right experiences to enable them to conform to the demands placed upon them. This will almost certainly involve a certain amount of formal "training." Although most dog owners know that they must train their dogs to make them socially acceptable, such a thought scarcely crosses cat owners' minds, or if it does, they reject it as only appropriate for "performing cats." Providing the right sort of experiences during the first few months of a kitten's life likely produces long-lasting effects, considering that this is the time when cats' personalities are forged, but more research is needed into precisely what those experiences should be.

The genetic basis for the cat's behavior must have changed as it adapted to living alongside man, even though the details are now lost in prehistory and ultimately impossible to trace. The average cat's personality is likely still changing, as some personality types fit modern conditions better than others. However, such haphazard change will not come about quickly enough to keep up with the pace of change that we now demand of our pets, as our own lifestyles change, so more direct intervention will be needed if the cat is to adapt at an acceptable pace.

We value our cats for their affectionate behavior, yet this trait has rarely been deliberately bred for-and then, only as an afterthought. This trait must have been accidentally selected for in the past, as loving cats get the best food!4 Nevertheless, "unfriendly" tendencies have persisted even among pet cats: in the 1980s experiments that defined the cat's socialization period, the experimenters noted "a small but constant percentage of the cats (about 15 percent) seem to have a temperament that is resistant to socialization."5 Even today, cats seem to possess a large range of variation in the genes that underpin temperament, learning, and behavior, providing the raw material for work on selective breeding for behavior and "personality," rather than just appearance.

As cat owners, we have various resources available to help individual cats adapt to today's crowded conditions, but many people seem unaware of many of these. Our challenge is to use these tools early in the cat's life, when it is still learning about its surroundings. The second and third months of life are the crucial period, during which the growing kitten learns how to interact socially-with other cats, with people, and with other household animals. As we have seen, it is during this period of its life that the cat learns both how to identify its social partners, and how to behave toward them in a way that will produce the desired outcome-a friendly tail-up, a lick behind the ear, a bowl of food, or a cuddle. More generally, this is also the time when cats learn how to cope with the unexpected-whether to be curious, accepting the risk of approaching and inspecting new things, or whether to play it safe and run away. Research shows that a cat's capacity to take risks can be subject to a strong genetic influence, but learning must also play a part.

A cat that has had only limited exposure to different kinds of people during its second and third months of life may become timid, retreating to its safe place whenever people it regards as unfamiliar come to the house. Without exposure to people before eight weeks of age a cat will become fearful of humans in general. However this is not the end of the process of socialization, just the necessary beginning: kittens must have the opportunity to make their own connections with different types of humans.

Many kittens homed at eight weeks may miss out on useful socialization experience with other kittens. The third month of life is when play with other kittens peaks, and feral kittens maintain strong social links with their peer group until they are about six months old. Veterinarians often (sensibly) advise that kittens should be kept indoors for the first few weeks after homing, to prevent them from straying: however, if there are no other cats in the house, they may miss out on a crucial phase in the development of their social skills.

Individual cats adopt different strategies when encountering the unfamiliar. Many withdraw, hide, or climb to a safe vantage point. A minority may become aggressive, perhaps those that have not been given the opportunity to retreat on previous occasions. Their owner may have run after them and scooped them up rather than allow them to withdraw. These cats quickly learn that the stress of an undesired encounter can be prevented by scratching, hissing, and biting. Some of these cats develop this tactic even further, preemptively striking out at people they don't know or who have previously been forced on them.

Cats also develop their own preferred tactics for dealing with other cats they do not know. When making their first social encounters as kittens, some simply flee; others attempt to stand their ground, and often get a swipe or worse for their efforts. Few attempt to engage in a friendly greeting, and even fewer find such a greeting reciprocated. Flight or fight thus often becomes the young cat's default response when meeting unfamiliar cats.

Owners who wish to add another cat to their household have an opportunity to manage the introduction to have a positive outcome for all concerned. We cannot take for granted that cats will immediately like one another. The new cat will likely feel stressed at being suddenly uprooted from its familiar surroundings and dropped into what it perceives as another cat's territory, and the resident cat will probably resent the intrusion. Therefore, it is usually best to start by keeping the new cat in a part of the house that the resident cat rarely uses, allowing it to establish a small "territory" of its own and getting to know its new owners before facing the challenge of meeting the resident cat face to face. The two cats will undoubtedly be aware of each other's presence, if only by smell, but this will be less stressful at this early stage than being able to see each other. Owners can build up some degree of familiarity between the two cats before a meeting takes place by periodically taking toys and bedding from each of the two cats and introducing them to the other while rewarding that cat with food treats or a game. This builds up a positive emotional link with the other cat's odor. The actual introduction should wait until after both cats no longer show any adverse reaction to the other's smell, and should be carried out in stages, starting with allowing the cats to be together for just a few minutes.6 Because cats carry a reputation for being untrainable, most owners are unaware that training can reduce the stress that cats can feel in situations where they would much rather run away. For example, owners can use clicker training (see chapter 6) to entice a cat to walk into its cat carrier, rather than forcing it in.7 Similar training could help cats overcome their initial fear of other potentially stressful situations-for example, encounters with new people. In general, cats need persuasion, not force, if they are to adopt a calm approach to new situations. If more owners understood the value of training, a great deal of stress could be avoided-for the cats, certainly, but also for the owner, if the cat's stress results in deposits of urine or feces around the house.

Training can also help cats to adapt to indoor living. Training a cat is a one-on-one activity that is mentally stimulating for the cat, and that also enhances the bond between cat and owner. It may also be useful in reducing some of the negatives of keeping a cat indoors. Cats need to express their natural behavior, and many owners understandably object to the damage to household furnishings that their cats unwittingly cause, for instance when sharpening their claws. In some countries, veterinarians remove the cat's claws surgically, but this may not be in the cat's best interest, and in some countries this intervention is illegal (see box on page 267, "Declawing"). A cat without claws may not only experience phantom pains from its missing toes, but is also unable to defend itself if attacked by other cats. Training the cat to claw only in specific places is a far more humane and straightforward alternative, especially if the cat has not yet developed a preference for soft furnishings.8 Although untested so far, training might also be useful in reducing cats' desire to hunt-or at least curtail their effectiveness as predators of wildlife. We know that when cats appear to be playing with toys, they think they're hunting, but we have no information on whether playing in this way reduces-or, just conceivably, enhances-their desire to hunt for real. If play does have an effect on this desire, how long does the effect last? Would a daily "hunting" game between owner and cat save the lives of garden birds and mammals? Is it possible to train a cat to inhibit its pounce?

We also know little about how experience affects the hunting habit in general. Cats vary enormously in how keen they are to go out hunting. That the basis for this is mainly genetic is unlikely, since only a few generations have elapsed since all cats had to hunt to obtain the right kind of food. Anecdotally, one of the arguments for allowing a female house cat to have one litter is that this is usually born in the spring, distracting the cat (provided its owners feed it well) from going out hunting and thereby learning its trade. Is there a "sensitive period" for perfecting predatory skills, after which the desire to hunt is unlikely to develop fully? Further study of this might not only save animals' lives, but also a great deal of aggravation between cat and wildlife enthusiasts.

Declawing Cats instinctively scratch objects with their front claws. Perhaps they do this to leave behind an odor or a visible sign of their presence, to alert other cats. They may also scratch because their claws are itchy: periodically, the whole of the outside of the claw detaches, revealing a new, sharp claw within. If this is not shed, maybe because the cat is arthritic and finds scratching painful, the whole claw may overgrow and cause the pad to become infected.

Some owners who object to scratch marks on their furniture seek to have the cat's front claws (and occasionally, too, its back claws) removed. Few veterinary procedures excite as much controversy as declawing (known technically as onychectomy). This is regarded as routine in the United States and the Far East, but is illegal in many places, including the European Union, Brazil, and Australia.

Declawing is a surgical procedure that involves amputation of the first joint of the cat's toes. The initial pain resulting from the procedure may be controlled with a.n.a.lgesics, but we do not know whether cats subsequently feel phantom pain due to the nerves that have been severed. However, cats and humans have nearly identical mechanisms for feeling pain, and four out of five people who have fingers amputated have phantom pain, so cats likely do as well. (I myself experienced phantom pain for more than ten years after most of the nerves in one fingertip were severed in an accident. I learned to ignore the pain because I knew it was meaningless-something cats are unlikely to be able to do.) Declawed cats are more likely to urinate outside their litter boxes than other indoor cats, possibly because of the stress of this phantom pain.

Claws are an essential defense mechanism for cats. While owners of indoor cats will argue that their cat never meets other cats, and so should never need their claws, a declawed cat that is picked up roughly by a person may resort to biting, unable to scratch to indicate its discomfort, and thereby cause a much more significant wound.9 Today's cats find themselves in a delicate situation. On one hand, they must adapt to meet our changing needs; on the other, they have a reputation for being a pet that is easy to maintain. Persuading many cat owners to train their cats, to spend that extra time and effort to change their pets' behavior, may be difficult. As such, we must focus on a genetic shift as well, taking the cat further down the road toward full domestication.

Ideally, cats that are predisposed to adapt to modern living conditions-to achieve the three goals outlined above-should be identified and then prioritized for breeding. This will not be entirely straightforward, since cats' personalities continue to develop after the normal age of neutering, so before any such breeding program can begin, research will be needed to separate the effects of the desired genes from those of the cat's social environment. Moreover, there is probably no single "perfect" set of genes that will enable cats to fit all the lifestyles that humankind will demand of them. The ideal indoor cat will almost certainly be genetically distinct from the ideal outdoor cat, since, among other differences, the owner can have far more influence on relationships with other cats if the cat is confined indoors.

We have three potential sources for such genes: house cats, pedigree cats, and hybrids. Conventional pedigree cats have been produced almost exclusively for their looks, not their behavior, so they are unlikely to be a rich source of new behavior traits.10 Pedigree cats are derived entirely from cats that have only ever had two functions: to hunt vermin and to be good companions. In most breeds, there seems to have been little direct selection for behavior, only looks. There are however a few interesting exceptions.

The Ragdoll is a semi-longhaired breed that was originally named for its extremely placid temperament. The first examples to be exhibited, bred in the early 1990s, went limp when picked up, almost as if the scruffing reflex was triggered by touch anywhere on the body, not just the back of the neck. It was once rumored that these cats were insensitive to pain, and animal welfare organizations raised concerns that people might be tempted to toss these cats around like cushions. The breed no longer shows such extreme behavior, but is still renowned for its easygoing temperament. A similar breed derived from the same original stock, the Ragam.u.f.fin, is described thus: "The only extreme allowed in this breed is its friendly, sociable and intelligent nature. These cats love people and are extremely affectionate."11 The genetic basis for these cats' relaxed sociality, at least toward people, is unknown, but could potentially be transferred to other cats by crossbreeding. Unfortunately for their welfare, ragdoll-type cats are reputedly vulnerable to attack from neighborhood cats, perhaps because they are simply too trusting, and for this reason many breeders advise prospective owners to keep them indoors.

Hybrids, crosses between Felis catus and other species, while initially produced mainly for their "exotic" appearance, have brought new genetic material into the domain of the domestic cat. Their behavior is often quite distinctive, so hybrids may potentially provide a source of genes that influence behavior that are not currently found in ordinary domestic cats.

The most widespread of these hybrids, the Bengal cat, may not offer any solution to the adapting the domestic cat to the twenty-first century, since its personality appears to have headed back toward that of its wild ancestors. The Bengal is a hybrid between domestic cats and the Asian leopard cat Prionailurus bengalensis. The latter species is separated by more than 6 million years of evolution from the domestic cat, and has never been domesticated in its own right; therefore, it would seem an unlikely starting point for a new breed of cat, were it not for its attractive spotted coat (referred to as "rosettes"). Domestic cats and Asian leopard cats will mate with one another if given no other option, but the resulting offspring are essentially untamable. During the 1970s, repeated breeding between these hybrids and domestic cats produced some offspring that retained the leopard cat's spots on the back and flanks, creating the current Bengal "breed."

Unfortunately, many Bengals possess not only wild-type coats, but also wild-type behavior, as this information from the Bengal Cat Rescue website confirms: This breed has a strong and sometimes dominant personality and although affectionate, lots are not simple lap cats. They can respond aggressively to discipline and to being handled . . . Their commonest problems are aggression and spraying. . . . Also hardly a week goes by when someone doesn't contact me about having bought or adopted a pair that are trying to kill each other. . . . Bengals enjoy climbing and this includes your clothes and curtains. They like exploring and are no respecter of ornaments or photographs. Often cat-aggressive, many will terrorise not just their own household but can actively seek out neighbors' cats and enter their homes to hurt them. They are not playing, they mean it.12 From the perspective of producing a docile pet, the Asian leopard cat was never going to be a good candidate for hybridization. This species is one of the few small wild cats that is not threatened with extinction; nevertheless, many zoos keep one or two specimens. These animals are, however, virtually untamable: zookeepers report that they are impossible to approach, let alone touch.13 From the perspective of seeking genes that could be useful for changing the domestic cat, some of the smaller South American cats could be better candidates for hybridization. In particular, Geoffroy's cat, similar in size to the domestic cat, and the slightly larger margay, are often friendly to their keepers when kept in zoos, and might therefore provide genetic material useful to the continued evolution of the domestic cat. The South American cats lost one pair of chromosomes soon after they diverged from the rest of the cat family some eight million years ago; this should mean that the offspring of a domestic cat and a South American cat is sterile, but surprisingly, hybrids with Geoffroy's cat can be fertile. The resulting "breed," usually known as the "Safari," was first created in the 1970s, but is still rare: it is usually still produced by mating the two original species, and the kittens, which grow into extra-large cats, fetch thousands of dollars. Breeding between these first crosses and ordinary domestic cats, the method used to produce the Bengal, seems to produce fertile offspring, but breeders apparently chose not to pursue this option. Hybrids with the margay, once referred to as the "Bristol," suffer from fertility problems and are apparently no longer bred. The margay is a tree-dwelling cat with double-jointed ankles, enabling it to climb down tree trunks as easily as other cats climb upward, to hang one-footed from branches like a monkey, and to leap twelve feet from one perch to another: such agility, if pa.s.sed onto its hybrids, might be appealing, but also excessive for most domestic purposes.

Bengal cat (above) and Safari cat (beneath) Several other types of cat have been produced through hybridization with other felids, but all have been bred for their "wild" looks, and none seem to offer more than curiosity value to the domestic cat's genome. These include the Chausie, a domestic cat crossed with the jungle cat Felis chaus, and the Savannah (serval Leptailurus serval), as well as many other oddities of doubtful provenance. Some are cla.s.sified as wild animals rather than as pets, much as wolf-dog hybrids are.14 These various hybrid breeds appear more of a side issue than a potential source of new genetic material to enrich the genome of Felis catus. The most promising of the species in behavioral terms, the more docile of the South American cats, are genetically incompatible with the domestic cat. The Old-World cats that are better matched genetically produce hybrids that are wilder, not calmer, than today's alley cats, so have little to offer.

Existing variation within Felis catus seems to be the best starting point for the completion of the cat's domestication. Plenty of modern cats combine an easygoing nature with a disinclination to hunt. Research has not yet indicated precisely how much of this variation is underpinned by genetics, but a significant proportion must be. Our goals should be to identify those individual cats with the best temperaments, and to ensure that their progeny are available to become tomorrow's pets.

One potential source of genetic variation has only recently emerged. Although house cats the world over are superficially similar, their DNA reveals that they are genetically distinct-as different under the skin as, say, a Siamese and a Persian are on the surface. Interbreeding between, for example, ordinary pet cats from China with their counterparts from the UK or the United States might produce some novel temperaments, some of which might be better suited to indoor life, or be more sociable, than any of today's cats.

Alley cats from the Far East (left) and Western Europe (right) Selection for the right temperament among house cats requires deliberate intervention; natural evolutionary processes, which have served the cat well so far, will not be enough. One obstacle is the increasingly widespread practice of neutering cats before they breed. With so many unwanted cats euthanized every year, arguing against the widespread use of this procedure is difficult. Still, if taken too far, widespread neutering likely favors unfriendly cats over friendly. Encouraging owners not to allow their cats to produce any offspring whatsoever removes all the genes that those cats carry from being pa.s.sed on to the next generation. Some of those genes have contributed to making those cats into valued pets.

When almost every pet house cat has been neutered-a situation that already applies in some parts of the UK-then we must fear for the next generations of cats. These will then mainly be the offspring of those that live on the fringes of human society-feral males, stray females, as well as those female cats owned by people who either do not care whether their cat is neutered or not, or have a moral objection to neutering. The qualities that allow most such cats to thrive and produce offspring are, unfortunately, those same behaviors we want to eliminate: wariness of people and effectiveness at self-sustaining hunting.

These cats will undoubtedly adapt their behavior to cope with whatever situations they find themselves in, but they are also likely to be genetically slightly "wilder" than the average pet cat-therefore, distinctly different from the "ideal" pet cat. Initially, the difference will probably be small, since some of the breeding cats will be strays that are genetically similar to pet cats. But as the decades pa.s.s, as fewer and fewer reproductively intact cats are available to stray, most of the kittens born each year will come from a long line of semi-wild cats-since these are the only cats that are able to breed freely. Thus, the widespread adoption of early neutering by the most responsible cat owners risks pushing the domestic cat's genetics back gradually toward the wild, away from their current domesticated state.

A study that I conducted in 1999 suggests that such extrapolation cannot be dismissed as science fiction.15 In one area of Southampton (UK), we found that more than 98 percent of pet cat population had been neutered. So few kittens were being born that potential cat owners had to travel outside the city to obtain their cats. This situation had clearly existed for some time: from talking to the owners of the older cats, we calculated that the cat population in that area had last been self-sustaining some ten years previously, in the late 1980s. We located ten female pets in the area that were still being allowed to breed and tested the temperament of their kittens after homing, when the kittens were six months old. Our hypothesis was that feral males must have fathered many of these kittens, since so few intact males were being kept as pets in the area, and all of these were young and unlikely to compete effectively with the more wily ferals. We found that on average, the kittens in those ten litters were much less willing to settle on their owners' laps than kittens born in another area of the city that still had a significant number of undoctored pet tomcats. There was no systematic difference in the way these two groups of kittens had been socialized, and the mother cats in the two areas were indistinguishable in temperament. We therefore deduced that even if only one of the two parents comes from a long line of ferals, the kittens will be less easy to socialize than if both parents are pets. The study was too small to draw any firm conclusions, but in the years since it was carried out, blanket neutering has become more widespread, and so the c.u.mulative effects of this on the temperament of kittens should be becoming more obvious.

Neutering is an extremely powerful selection pressure, the effects of which have been given little consideration. At present, it is the only humane way of ensuring that there are as few unwanted cats as possible, and it is unlikely ever to become so widely adopted that the house cat population begins to shrink. However, over time it will likely have unintended consequences. Consider a hypothetical situation. A century or more ago, when feline surgery was still crude, society generally accepted that most cats would reproduce. Imagine that a highly contagious parasite had appeared that sterilized cats of both s.e.xes while they were still kittens, but otherwise left them unaffected, so that they lived as long as an unaffected cat. Any cats that happened to be resistant to that parasite would be the only cats able to breed-so, within a few years the parasite, deprived of susceptible hosts, would die out.

The only significant difference between such a hypothetical parasite and neutering is that the latter does not require a host (a cat) to continue: it lives as an idea, and so is detached from its effects.16 Because neutering inevitably targets those cats that are being best cared for, it must logically hand the reproductive advantage to those cats that are least attached to people, many of which are genetically predisposed to remain unsocialized. We must consider the long-term effects of neutering carefully: for example, it might be better for the cats of the future as a whole if neutering programs were targeted more at ferals, which are both the unfriendliest cats and also those most likely to damage wildlife populations.

We need a fresh approach to cat breeding. Pedigree cats are bred largely for looks, not with the primary goal of ensuring an optimal temperament-although an adequate temperament is, of course, taken into account in the majority of breeds. Random-bred cats are under siege from neutering; even if this widespread practice is not making each successive generation a little wilder than the previous, it is highly unlikely to be having the opposite effect. So while cats' looks and welfare both have their champions, the cat's future has none.

Then again, why should it? Cats have always outnumbered potential owners. Why should that situation change? Cats have become more popular, not less, so there should be more homes available for them, not fewer, than there were a few decades ago. Apart from the (significant) minority of cat-haters, the general public is more tolerant of cats than of dogs. We cannot guarantee, however, that these apparent givens will continue.

Recent decades have witnessed immense changes to the way dogs are kept, especially in urban areas, with a proliferation of "clean up after your dog" regulations, no-dog parks, and legislation aimed at protecting the public from dog bites. We expect dogs to behave in a much more controlled and civilized way than they did half a century ago. Are similar restrictions on cat-keeping just around the corner? Will gardeners and wildlife enthusiasts unite to produce legislation that restricts cats to their owners' property? If such pressures do appear, they will be easier to head off if cat-lovers are already taking steps toward producing a more socially acceptable cat. At the same time, the cats themselves will benefit if they find it easier to cope with the vagaries of their pets' social lives.

Ultimately, the future of cats lies in the hands of those who breed them-not those whose eye is primarily on success in the show ring, but those who can be persuaded that an improved temperament, not good looks, should be the goal. The genetic material is available, although more science is needed to devise the temperament tests that will locate which individual cats carry it; many cats that appear well-adapted to life with people will have received an optimal upbringing, rather than being anything special genetically. The relevant genes are probably scattered all over the globe, so ideally we need collaboration between cat enthusiasts in different parts of the world.

Such human-friendly cats, however cute, are unlikely ever to command much of a price. Expectations that non-pedigree kittens should be free, or nearly so, will take a long time to die out. Commercial breeding of non-pedigree cats may never be viable. Well-adjusted kittens need a wealth of early experience that even some pedigree breeders struggle to provide. Providing this level of care is cost-effective only if kittens are bred in people's homes, the very type of environment into which they will move when they become pets.

Meanwhile, the way that cats are socialized has much room for improvement. Both breeders and owners can play a part in this, since kittens adapt to their surroundings throughout their second and third months of life. In this context, the continuing policy of some cat breeders' a.s.sociations to prohibit homing until a kitten is twelve weeks of age demands careful scrutiny: it may provide extra socialization with littermates, but often at the cost of learning about different kinds of people, and the development of a robust strategy for dealing with the unfamiliar. For adult cats, training, both in the general sense of providing the right learning experiences as well as teaching them how to behave calmly in specific situations, could improve each cat's lot considerably, if only its value was more widely appreciated.

Finally, we must continue to research why some cats are strongly motivated to hunt, while the majority are content to doze in their beds. Science has not yet revealed to what extent such differences are due to early experience, and how much to genetics, but ultimately it should be possible to breed cats that are unlikely to feel the need to become predators, now that we can easily provide them with all the nutrition they need.

Cats need our understanding-both as individual animals that need our help to adjust to our ever-increasing demands, and also as a species that is still in transition between the wild and the truly domestic. If we can agree to support them in both these ways, cats will be a.s.sured a future in which they are not only popular and populous, but are also more relaxed, and affectionate, than they are today.

Further Reading

Most of my source material for this book has consisted of papers in academic journals, which are often difficult to access for those without a university affiliation. I've included references to the most important of these in the notes, with Web links if they appear to be in the public domain. For those readers who wish to take their study of cats further without first requiring a degree in biology, I can recommend the following books, most of them written by knowledgeable academics but with a more general audience in mind.

The Domestic Cat: the Biology of Its Behaviour, edited by Professors Dennis Turner and Patrick Bateson, is now available in three editions; all are published by Cambridge University Press, the most recent in 2013. These books consist of chapters written by experts in different aspects of cat behavior.

My own The Behaviour of the Domestic Cat, 2nd edition (Wallingford, UK: CAB International, 2012), coauth.o.r.ed by Drs. Sarah L. Brown and Rachel Casey, provides an integrated introduction to the science of cat behavior, aimed at an advanced undergraduate audience. Feline Behavior: A Guide for Veterinarians by Bonnie V. Beaver (St. Louis, MO: Saunders, 2003) is, as its t.i.tle indicates, aimed at veterinary surgeons and veterinary students.

For the various stages in the history of the cat's life with humankind, Jaromir Malek's The Cat in Ancient Egypt (London: British Museum Press, 2006), Donald Engel's Cla.s.sical Cats (London: Routledge, 1999), and Carl Van Vechten's The Tiger in the House (New York: New York Review of Books, 2006) provide specialist accounts.

Carrots and Sticks: Principles of Animal Training (Cambridge: Cambridge University Press, 2008) by Professors Paul McGreevy and Bob Boakes from the University of Sydney, Australia, is a fascinating book of two halves: the first explains learning theory in accessible language, and the second contains fifty case histories of animals (including cats) trained for specific purposes, ranging from film work to bomb detection, each ill.u.s.trated with color photographs of the animals and how they were trained.

For cat owners seeking guidance on a problem cat, there is often no subst.i.tute for a one-to-one consultation with a genuine expert, but these can be hard to find. The advice given in books by Sarah Heath, Vicky Halls, or Pam Johnson-Bennett may be helpful. Celia Haddon's books may also provide some light relief.

Notes.

All Web addresses mentioned in the Notes are active as of April 2013.

Introduction.

1. This ratio takes many millions of unowned animals into account, and also incorporates a guess as to the numbers in Muslim countries where dogs are rare.

2. The Prophet Muhammad is said to have loved his cat Muezza so much that "he would do without his cloak rather than disturb one that was sleeping on it." Minou Reeves, Muhammad in Europe (New York: NYU Press, 2000), 52.

3. Rose M. Perrine and Hannah L. Osbourne, "Personality Characteristics of Dog and Cat Persons." Anthrozos: A Multidisciplinary Journal of the Interactions of People & Animals 11 (1998): 3340.

4. A recognized medical condition, referred to as "ailurophobia."

5. David A. Jessup, "The Welfare of Feral Cats and Wildlife." Journal of the American Veterinary Medical a.s.sociation 225 (2004): 137783; available online at www.avma.org/News/Journals/Collections/Doc.u.ments/javma_225_9_1377.pdf.

6. The People's Dispensary for Sick Animals, "The State of Our Pet Nation . . . : The PDSA Animal Wellbeing (PAW) Report 2011." Shropshire, UK: 2011; available online at tinyurl.com/b4jgzjk. Dogs scored a little better for social and physical environments (71 percent) but worse for behavior (55 percent).

7. The situation for pedigree dogs in the UK has been summarized in several expert reports, including those commissioned by the Royal Society for the Prevention of Cruelty to Animals (www.rspca.org.uk/allaboutanimals/pets/dogs/health/pedigreedogs/report), the a.s.sociate Parliamentary Group for Animal Welfare (www.apgaw.org/images/stories/PDFs/Dog-Breeding-Report-2012.pdf), and the UK Kennel Club in partnership with the re-homing charity DogsTrust (breedinginquiry.files.wordpress.com/2010/01/final-dog-inquiry-120110.pdf).

Chapter 1.

1. Darcy F. Morey, Dogs: Domestication and the Development of a Social Bond (New York: Cambridge University Press, 2010).

2. Quoted in C. A. W. Guggisberg, Wild Cats of the World (New York: Taplinger, 1975), 3334.

3. These cats are now extinct on Cyprus, displaced by the red fox, another introduction, which is now the only land-based carnivorous mammal on the island.

4. For a more detailed account of these migrations, see Stephen O'Brien and Warren Johnson's "The Evolution of Cats," Scientific American 297 (2007): 6875.

5. The spelling lybica should more correctly be libyca, "from Libya," but most modern accounts use the original (incorrect) version.

6. These "lake dwellers" built villages on sites that now lie beneath the margins of lakes, but were probably fertile dry land at the time.

7. Frances Pitt (see note 9 below) claimed that the Scottish wildcat would have joined its English and Welsh counterparts in extinction, had it not been for the call-up of the younger gamekeepers to fight in the Great War.

8. Carlos A. Driscoll, Juliet Clutton-Brock, Andrew C. Kitchener, and Stephen J. O'Brien, "The Taming of the Cat," Scientific American 300 (2009): 6875; available online at tinyurl.com/akxyn9c.

9. From The Romance of Nature: Wild Life of the British Isles in Picture and Story, vol. 2, ed. Frances Pitt (London: Country Life Press, 1936). Pitt (18881964) was a pioneering wildlife photographer who lived near Bridgnorth in Shropshire.

10. Mike Tomkies, My Wilderness Wildcats (London: Macdonald and Jane's, 1977).

11. This and the following two quotations are from Reay H. N. Smithers's "Cat of the Pharaohs: The African Wild Cat from Past to Present," Animal Kingdom 61 (1968): 1623.

12. Charlotte Cameron-Beaumont, Sarah E. Lowe, and John W. S. Bradshaw, "Evidence Suggesting Preadaptation to Domestication throughout the Small Felidae," Biological Journal of the Linnean Society 75 (2002): 36166; available online at www.neiu.edu/~jkasmer/Biol498R/Readings/essay1-06.pdf. In this paper, which came before Carlos Driscoll's DNA study making cafra a separate subspecies, the Southern African cats are listed as Felis silvestris libyca.

13. Carlos Driscoll et al., "The Near Eastern Origin of Cat Domestication," Science (Washington) 317 (2007): 51923; available online at www.mobot.org/plantscience/resbot/Repr/Add/DomesticCat_Driscoll2007.pdf. The data discussed can be found in the online Supplemental Information, Figure S1.

14. David Macdonald, Orin Courtenay, Scott Forbes, and Paul Honess, "African Wildcats in Saudi Arabia" in The Wild CRU Review: The Tenth Anniversary Report of the Wildlife Conservation Research Unit at Oxford University, ed. David Macdonald and Francoise Tattersall (Oxford: University of Oxford Department of Zoology, 1996), 42.

15. The estimate of fifteen to twenty comes from Carlos Driscoll of the Laboratory of Genomic Diversity at the National Cancer Inst.i.tute in Frederick, Maryland, who is currently working to pinpoint where these genes lie on the cat's chromosomes, and how they may work.

16. See note 13 above.

17. O. Bar-Yosef, "Pleistocene Connexions between Africa and Southwest Asia: An Archaeological Perspective," The African Archaeological Review 5 (1987), 2938.

18. Carlos Driscoll and his colleagues have discovered five distinct types of mitochondrial DNA in today's domestic cats; mitochondrial DNA is inherited only through the maternal line. The common maternal ancestor of these five cats lived about 130,000 years ago; over the next 120,000 years, her descendants gradually moved around the Middle East and North Africa, their mitochondrial DNA mutating slightly from time, before a few of them happened to become the ancestors of today's pet cats.

Chapter 2.

1. J.-D. Vigne, J. Guilane, K. Debue, L. Haye, and P. Gerard, "Early Taming of the Cat in Cyprus," Science 304 (2004): 259.