Origins of the cat

Although the ancestors of the domestic cat (Felis silvestris catus) first appeared about 35 million years ago (MYA) during the late Eocene, the cat family or Felidae, to which all living cat species belong, emerged somewhat later during the Miocene about 10–11 MYA (Johnson & O’Brien, 1997; O’Brien et al., 2008). Morphological and molecular studies of phylogenetic relationships among living felids indicate that the 37 extant species can be divided up into 8 major phylogenetic groups or lineages: the Panthera lineage; the bay cat lineage; the leopard cat lineage; the caracal lineage; the ocelot, lynx and puma lineages; and, finally, the domestic cat or Felis lineage (Leyhausen, 1979; Collier & O’Brien, 1985; Salles, 1992; Johnson & O’Brien, 1997; O’Brien et al., 2008). The latter is believed to have diverged from the others around 6.2 MYA, and comprises four species of small cats that seem to have originated around the Mediterranean basin: the jungle cat (Felis chaus), the black-footed cat (F. nigripes), the sand cat (F. margarita) and the wildcat (F. silvestris) (O’Brien et al., 2008).

Introduction

Research activities on the welfare of domestic cats have increased greatly in recent years. Of particular note is the emergence, especially in the USA and to a lesser extent in the UK, of shelter medicine as a veterinary specialist discipline in its own right. Its emergence is immensely gratifying, and is already leading to major improvements and refinements in the ways that overpopulation, one of the most important global issues in cat welfare, is being tackled. Significant advances are being made in the ways shelters are managed and cats are cared for. In addition, we are beginning to understand better why cats end up in shelters, what to do about it and how to promote successful adoptions. As a result, the number of healthy cats and kittens euthanised in shelters has decreased in some countries, although it still remains much too high.

There have also been improvements in our understanding of the needs of cats and how they can be met, whether cats are housed in the home, the shelter or boarding cattery, the veterinary surgery or the research facility. Innovative ways of enriching the environment of cats in order to meet these needs and improve their welfare are being developed and applied in practice. By enriching the lives of cats under our care, we also enrich our own lives.

Introduction

The whole concept of different breeds of cat is a relatively modern one extending back only about 150 years. Other domesticated animals have a breed history much older; dogs and horses in particular were deliberately domesticated to ‘serve’ humans and were, to some extent, selectively bred from very early in their relationship with people to fit their needs. Thus the idea of selecting for different physical ‘types’, whether size, mass, speed, or certain mental abilities or characteristics is something human beings have pursued, albeit in a somewhat random way, probably for several millennia. By the eighteenth century this practice in some domestic species was quite refined, with a large number of recognisably different types of dogs, horses and, following the agricultural revolution from the mid eighteenth century onwards, of various farm animals such as cattle, sheep, pigs and fowl, all developed to meet different needs in human society. Deliberate selection had not yet been applied to the domestic cat, but by the latter half of the nineteenth century there was a growing understanding of basic genetics and as a result people realised they could select for, perpetuate and even improve certain attractive or distinctive features in domestic animals. Mendel’s findings on the principles of heredity were initially ignored, but by the beginning of the twentieth century were gaining acceptance. It was discovered, without the reasons being completely understood, that by breeding like to like or mating together closely related animals, offspring could exhibit the features particularly valued and sought after. Although the domestic cat has had a relationship with humans for at least 4000 years, this relationship has principally been one of exploitative captive, where the cat provided an effective control of rodents and other vermin in both urban and rural environments. In performing these functions the cat was regarded as extremely useful and gradually also respected as a companion animal (see Chapter 7).

Introduction

When a cat behaves socially towards a human, the person has been treated as though he or she were a cat, although very possibly a particular type of cat. Anybody who loves cats is irresistibly drawn to treat them as though they had some of the characteristics of humans. Cat owners project themselves into the heads of cats and, in so doing, empathise with them. Whether or not they are right to do so raises a big and largely unanswered question. In any event, owners will have seen their cats behave in ways that are often puzzling and enigmatic. Why are they sometimes so friendly and at other times so distant? Those who study the behaviour of cats scientifically are often asked to provide the real answers to such questions. Unfortunately, many aspects of cat behaviour that most interest the lay public have not been the subject of extensive investigation. In part this is because the implied question is: ‘Why does the animal need to behave in this way?’ This is a question about the current utility to the animal of behaving in a particular fashion and is not an easy one to answer. The question also raises questions about the evolution of the domestic cat.

To understand the biological value to the animal of behaving in a particular fashion, scientists ask how the behaviour of a cat, freely living in a natural environment, helps it to survive and breed now. To understand the evolution of the behaviour, they must speculate about the natural environment of the modern cats’ wild ancestors. If some patterns of behaviour have worked to the individual’s advantage better than others in the past and they were inherited, they would eventually tend to be shared by most members of the cat population. The presumption is that by the process of Darwinian evolution, cats behave in a way that is well-adapted to the type of social and physical environment in which their ancestors lived.

Introduction

With so many people owning cats, understanding cat behaviour in the human home is important. Cats are the largest pet population in the USA. In 2011, of 374 million pets in the USA, 86.4 million were cats. In the same year, 78.2 million were dogs (ASPCA, 2012). In 2010 in the USA more than 22 billion dollars were spent on cat food (American Pet Products Association, 2011). Most of the research on domestic cat behaviour has been obtained from studies of cats in shelters, in laboratory situations or in free-roaming (feral) situations. Veterinarians and animal behaviourists have relied on information gained from studies of these populations for information about the normal behaviour of domestic cats. Most of the interactions people have with their cats is in the home. Domestic cats rarely accompany their owners outside of their homes. According to the recent American Pet Products Manufacturers Association’s Pet Owners Survey, 3% of cat owners in the USA take their pets with them when they travel for at least two nights compared with 19% of dog owners (American Pet Products Association, 2011). In a recent repeated-measures study, cats were present with their owners 6% and dogs were present with their owners 19% of the time the owners were outside of the home (Friedmann et al., 2010). However, very little is known about cats’ behaviour in the home.

Introduction

The almost ubiquitous presence of domestic cats in human society is evidence that people value this species highly, both as companions and as biological control of pests. In some countries, including the UK, USA and China, they have out-competed other species in becoming the most common household pet (Bernstein, 2005). However, the presence of cats can also give rise to a range of problems to humans and other species and they may experience welfare problems themselves. As cats can survive and reproduce successfully outside of human care, or indeed with human care but without human intent, their populations and the problems these present may increase to a level deemed unacceptable to society and hence the motivation for ‘cat population management’. The global number of domestic cats is a notoriously difficult figure to estimate; Jarvis (1990) suggested 400 million cats globally; however, an unpublished report to the World Society for the Protection of Animals (WSPA) in 2007 used published data on pet statistics and questionnaires of non-governmental organisations and veterinary surgeons to gather information from 194 countries, leading to an estimate of 272 million cats worldwide of which 58% were thought to be ‘stray’ or ‘feral’ (see the discussion that follows regarding these terms).

When going through the adoption process for a new puppy there is typically a focus on breed identification with an eye on what to expect when the dog grows up in terms of behaviour as well as body shape and size. Talk to someone who is thinking of adopting a kitten and typically little attention is given to breed or breed composition. Instead, thought is given to hair coat length, colour and pattern along with a concern about an appropriate place from which to adopt a kitten (Karsh & Turner, 1988).

The overwhelming practice in selecting a kitten is to adopt a generic type, commonly referred to as a domestic shorthair (DSH) or longhair (DLH). It appears as though not much thought is given to the behavioural profile of the cat as an adult. The logic seems to be ‘a cat is just a cat’. In other words, you get what you get. Contrast that with someone who adopts a Jack Russell Terrier dog knowing that the behaviour – reactive and snappy – will be quite different from the Golden Retriever they had before.

Puppy adopters commonly select a breed based on what they expect in the way of behaviour as the puppy grows up; knowing the breed, one can predict the future behaviour to some extent (Hart & Hart, 1985, 1988). While people adopting kittens do not seem to consider the future behaviour of the kitten, they are certainly affected by the behaviour of their cat as adulthood is reached. The behaviours that many cat caregivers find desirable include being affectionate towards the human family members and socially outgoing; and you can add in being good at litter box use. Behaviours that are universally undesirable are being aggressive towards human family members, overly fearful of visitors and urine marking in the house. What astute cat people realise is that some behavioural differences among cats are genetically based. And, when it comes to purebred cats, there are striking differences in behaviour – at least between some breeds – as there are in dogs.

Introduction

People and cats together

Domestic cats (Felis silvestris catus) are among the most common companion animals. This is particularly true in cultures with an Islamic background, where dogs are less acceptable as companion animals than in Western societies (Chapter 8). In Austria, for example, a human population of 8 million owns more than 2 million cats, in contrast to just about 700,000 dogs (Kotrschal et al., 2004). Consequently, the behaviour of cats and their interaction with people has attracted scientific interest (Leyhausen, 1960; Turner, 1991; Bradshaw, 1992; Turner & Bateson, 2000). Particularly in rural settings, the association between cats and people may still be loose; cats are mainly tolerated as pest controllers, but often people also sympathise with these cats and feed them. In urban areas most cats, nowadays, tend to be social companions of their owners. In parallel with the increase of one-person households, the keeping of companion cats seems on the rise. In order to ensure their safety, many of these urban cats are kept indoors. Such indoor cats tend to interact more with, and are more ‘attached’ to their owners than cats that have the option of going outdoors (Stammbach & Turner, 1999). People often engage in close and long-term relationships with their cats, and owners and cats may develop complex idiosyncratic and time-structured interactions (Wedl et al., 2011).

Introduction

Many previous accounts of communication between domestic cats have been largely based on a traditional ethological approach. The signals and the context in which they occur have been described and related to the kind of environment signaller and receiver can expect to find themselves in, and to the sensory capabilities of the receiver. For example, this approach explains the use of scent signals by domestic cats as products of both their acute sense of smell, which probably evolved primarily in relation to detection of food, and also their origin as territorial animals which needed to communicate with neighbours that they might rarely encounter face to face. However, the domestic cat is the product of two distinctly different phases of evolution, the first as a wild, largely solitary, predator, and the second as a commensal and then semi-domesticated social species, living in an increasingly dependent relationship with humans. There are few studies of the communicative repertoire of the ancestral species, Felis silvestris libyca, and it is now clear from the distribution of libyca DNA that many wild cats, including those from Africa and the Middle East, are, in varying proportions, hybrids between wild F. silvestris subspecies and domesticated F. s. catus (Driscoll et al., 2007). Commensalism will have brought with it new selection pressures on communication, largely intraspecific and resulting from the higher density at which these cats live, by comparison with that of their solitary ancestors.

Of all domestic animals, cats are the most capable of breeding, giving birth to offspring and raising their young without human care or intervention. In the classic example, a mother cat that has secretly gone through parturition in some obscure location reveals a litter of perfectly healthy kittens to the human family members after the kittens have been born. This romanticised view of feline motherhood has given way to the more formalised cattery operation, or the family breeder where a mother cat may give birth to her litter of kittens in the midst of an overly concerned family audience.

In this chapter, following some general information about parturition and the care of newborn, normal maternal behaviour is discussed along with comments about problems with various aspects of maternal behaviour. Problems with maternal behaviour manifest themselves primarily as either lack of proper attention to the kittens, resulting in inadequate care and nutrition, or cannibalism of the kittens. Of course, a necessary aspect of reproduction is the mating, which in this chapter is dealt with mostly in the context of intentional mating of females with selected males, in a home or cat-breeding facility.

Introduction

This chapter reflects an amazing ‘success story’, the story of a predator species, the domestic cat, which has more or less conquered the world within a few thousand years – partly with the help of humans, but mostly because of its amazing flexibility. It is not a new story and therefore easy to summarise. The summary is based upon three chapters in the second edition of this book, namely Macdonald et al. (2000), Liberg et al. (2000), and Fitzgerald and Turner (2000), but of course updated with later findings. Nor is that success story without ecological consequences which continue to fire the debate between cat friends and cat foes – or at least cat lovers and conservationists. Both sides of this debate should view the evidence before making hasty judgements.

Solitary life versus group-living: a question of resource availability

The domestic cat as a species and quite possibly as an individual shows amazing flexibility in its sociality toward conspecifics. Its ancestor, the North African wildcat, F. s. libyca, was (and is) indeed a solitary, territorial species, which presumably made use of the rodent populations concentrated in and around grain storage facilities of early farming settlements (see Chapter 7). As the likely story goes, this was beneficial to the farmers and they began provisioning the ‘wild’ cats with extra food, their home ranges became concentrated – and overlapping – around these human settlements and storage facilities, representing the first step toward domestication. The Resource Dispersion Hypothesis proposes that the dispersion of resources may be such that the smallest territory providing adequate security for the primary social unit (mother and offspring) may also support additional group members (Macdonald et al., 2000).