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Chapter 2 Development of companion animals.

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A cat is a lion in a jungle of small bushes.

--Indian Proverb

KEY TERMS

domestication

phenotypes

socialization

traits

feral

trap-neuter-release (TNR)

neoteny

breeds

evolution

selective breeding

genome

mitochondrial DNA (mtDNA)

nuclear DNA

purebred

tame

fancy

kennel club

affiliative

livestock

shelf pets

pocket pets

selfs

rare breeds

Companion animals are, in general, domesticated species. That is, they have been modified in various ways from some wild ancestor to make them more appropriate for living in close proximity with humans. Domestication is a process that takes place over a period of many generations of a species. It is important to recognize that species itself is a dynamic term that describes the relative state of a group of organisms in relation to other groups of organisms. Species are not static. A species is constantly responding to the environmental pressures and other species that surround it. It is on an evolutionary trajectory that will allow it to survive within its environment, or if the changes are too great or sudden, it will become extinct. Domestication occurs when humans alter that evolutionary trajectory for some part of a species' population, and instead of continuing to adapt to the natural environment where it has been found, it begins to adapt to an environment created by and associated with humans. This process is both biological and cultural (Clutton-Brock, 1992). As the species changes, or evolves, its place and role in relation to humans will change. It is an iterative process as changes in biology and behavior suit it to fill new niches with humans, and this in turn will suggest still other opportunities.

Domestication and its results are one of the most visible, and at the same time, poorly understood biological events to impact a species. All species have not been domesticated in the same way, and different species may be at different stages of the domestication process. Several key elements are generally common to all types of domestication. The first is that the population of animals (or plants for that matter) must show a range in phenotypes. This phenotypic variation must be correlated with underlying genetic variation. It is important that at least some members of the population are reasonably tolerant of humans being nearby, and through experience demonstrate a greater level of socialization to people. Most of the significant domesticated species were derived from wild ancestors that lived in social groups, whether herds, packs, or flocks. Dogs, horses, pigs, and chickens are just a few examples. Domestic cats are a rare exception. Other than lions, wild felines do not spend significant time in social groups. Contact with other members of the species is typically limited to mating, and mother-offspring interactions. Domestication has helped to make cats more tolerant of one another.

While individual members of a species may be tamed, allowing or tolerating a high degree of close human proximity or contact, it will retain instinctive behavior patterns, and will not directly pass its tameness on to it young. One of the first stages of domestication is the selection, whether or not intended, of individuals that are easiest to approach and socialize, and least likely to show instinctive patterns of attack or fleeing.

When young, many species will have a brief time frame when they will socialize to conspecifics. This sensitive period may be just several hours long, or extend for a number of weeks. Immediately after hatching, geese and ducks will imprint on the first moving object that they see (Immelman, 1972). They will then follow this object or individual, and use it as model for social bonding and orientation. This would usually be their parents and would ensure that their primary orientation is towards a conspecific. These birds are precocial, leaving the nest soon after hatching so it is critical that they develop a bond with their parents quickly after birth. Canines will generally give birth in some sort of den. The pups are helpless at birth and the mother provides extensive care to her young. They socialize gradually, first to their mother, then to siblings, and then to conspecifics that may be part of a pack or social group. Another aspect of domestication may be the lengthening of this socialization period. The critical period for dogs is quite long, lasting from about 3 to 19 weeks after birth. These expanded socialization windows make it easier to raise an animal that will be calm and accepting of humans, and other animals. Wild animals when taken as young may be tamed by socializing them with people. This is an uncertain process, however, and when the animal reaches sexual and physical maturity may revert to instinctive behaviors and present a danger to people and other animals.

A second change is the enhancement of the reproductive potential of the species. Domestication typically included an economic motivation that usually required the production of more individual animals. The reproduction of many wild species is modulated by seasons, social structure and status, the availability of prey or food, and other factors. Domestic species, when compared to wild counterparts, may begin breeding at a younger age, more frequently, have larger numbers of young, and be non-selective in the acceptance of mates, accepting the mate provided by humans.

Once a species is easy to handle and breed, it is possible to identify and select animals that exhibit desirable traits, and breed them for a specific purpose or appearance. The extent of variation that we may see in a domestic species will be a function of both the genetic structure and variation that was present in the original population of wild ancestors, and mutations that may be identified, preserved, and used in breeding programs. On the other hand, breeding animals within a small population can result in inbreeding and the loss of genetic variation and an increased probability of inherited diseases. Over time, accumulated differences between the wild and domestic forms may preclude hybridization, cross breeding, or the production of viable, fertile offspring.

There are times when domestic animals have escaped or otherwise returned to living on their own in a free-living or wild state. The mustangs of the American West are descendents of horses brought to the New World by explorers and settlers. Pigeons, common in large numbers in both urban and rural areas, are the prolific progeny of ancestors brought to America as a ready source of food in the early colonies (Figure 2-1). Over time, descendents of these domestic species living in such a fashion will begin to re-adapt to a free-living or wild existence. Descendents of a domestic species living in this way are called feral. Among the most common changes is a reacquired wariness toward people. This is likely a function of both limited or non-existent opportunities to socialize when young, and eventual genetic changes in the population. Unfortunately, one of the changes resulting from previous domestication that tends to persist is the increased reproductive capacity. As a result, a high rate of reproduction, often coupled with limited natural predators, may result in a population with unchecked growth that may threaten the local environment. Efforts to deal with feral cat populations had typically relied on lethal control methods. Trap-neuter-release (TNR) programs are now becoming a more common approach (Slater 2002). Feral cats are caught using live traps, sterilized, often vaccinated for rabies and other diseases, and then released back in the area where they were captured. Caretakers will then ensure that the cats are provided with food, and often some type of shelter in severe weather. This is a compromise approach. Feral cats are difficult to tame and it may not be possible to place them in new homes. At the same time there is limited public support for euthanizing cats that are generally healthy. In addition, lethal control has not been highly effective controlling populations of free-roaming cats. Sterilization helps to control population growth, and can result in a slow decline in numbers through attrition by natural causes.

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Another common result of domestication is neoteny, or maintaining juvenile physical and behavioral characteristics into maturity. Many breeds of dogs maintain the turn down/flop ears, rounded face, and shortened muzzles of puppies. Adult dogs may also exhibit the solicitation behaviors usually seen in puppies. Both wolf and dog puppies will lick the muzzle of adult dogs in greeting them. It is not uncommon for adult dogs to offer their owners "kisses" when they greet them, or when they want to solicit food or attention. Domestic cats will retain the predatory play behavior observed in kittens of wild felines. In wild kittens, this play will develop into highly efficient predatory sequences. In domestic cats this play may or may not develop into the ability to catch and kill prey. It will be strongly dependent on early experiences as a kitten (Fitzgerald & Turner, 2000; Leyhausen, 1979).

In summary, domestication will generally have the following impact on a species:

* Selection, direct or indirect, for tameness, or tolerance for the presence of people

* Changes in reproduction

** Earlier sexual maturity

** Breed more frequently

** Breed indiscriminately

** Larger numbers of young

** Neoteny of physical and behavioral traits

** Overall changes in size--usually smaller domesticated versions, though

"giants" may also be found

DOMESTICATION/DEVELOPMENT OF COMPANION ANIMALS

Dogs

Dogs are the most diverse single species on the planet. No other species shows the extraordinary range in size of the dogs. The size will range from the 0.5 kg Chihuahua to the 80 kg Great Dane. One could hardly imagine that the nearly hairless hide of the Chinese crested and the long, corded coat of the puli could come from two individuals of the same species (Figure 2-2). There are probably several hundred different breeds and types of dogs worldwide, 152 of which are recognized by the American Kennel Club (Crowely & Adelman, 1998), and in one way or another they tend to fall between these different extremes. The variation in dog behaviors may be nearly as great as their physical differences. Some dogs point, others retrieve, still others herd and some guard. The only constant may be that whatever the look and behavior, someone, somewhere liked it, wanted it, and chose to breed dogs in a way to either develop or maintain the trait.

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There has been a recent revolution in the study of dog evolution (Zawistowski & Patronek, 2004). New methods in molecular genetics have provided a vast new set of data revealing some unexpected surprises. When combined with traditional studies of fossils and archaeological evidence we have a very complicated story that we are only starting to understand. The dog has long been considered the first species domesticated by humans (Clutton-Brock, 1995). The wolf was thought to be the traditional ancestor of the dog. However, given the wide physical and behavioral variation seen in the dog, it was not uncommon to believe that other wild canids must have been involved in the development of the dog. Darwin studied a variety of domestic species and the role that selective breeding can play in the evolution of a species. He bred several varieties of domestic pigeons and frequently attended animal shows and talked to breeders. Based on his observations, Darwin did not believe that all dogs could have descended from a single ancestral species (Darwin, 1859/1979). Konrad Lorenz (1954) also believed that several different wild canids, including the jackal, gave rise to the many different breeds of dogs. Lorenz would later retract his statements regarding jackals and their role in dog evolution (Lorenz, 1975). He did believe that the northern, spitz breeds were those most closely related to wolves. In contrast to Darwin, who felt that the study of domestic animals could contribute to a better understanding of evolution and natural selection, Lorenz was among the modern scientists who disdained the value of studying domestic animals (Serpell, 1995). This attitude is changing, and the study of dog biology and behavior, combined with information on its genome may provide important insights into gene regulation and function (Ostrander & Kruglyak, 2000; Sutter & Ostrander, 2004).

Canine Archaeology. Archaeological evidence related to the origin of the dog can be difficult to interpret. Fossil bone fragments that came from a wolf or a dog may be very similar at the early stages of evolutionary separation. This is especially true in those areas where the wolf species of the time were small in size, closer to the size of the early dogs (Harrison, 1973; Olsen, 1985). There are also limited numbers of clearly identified dog fossils available for the earliest period when the separation from wolves may have been happening. At the same time, the animals giving rise to those fossils may have shown important behavioral differences. Since behavior does not fossilize, the context and locations in which remains are found are significant. Remains found in proximity to other human artifacts would suggest that the animal was associated with people. These data have in general placed the domestication of dogs in an era about 14,000 BP (Beneke, 1987; Clutton-Brock, 1995), or late Paleolithic period. An important find that also suggested a significant social relationship with dogs was of a puppy skeleton that had been buried with a human 12,000 BP (Davis & Valla, 1978). Found in the upper Jordan Valley of Israel, the canine skeleton was clearly that of dog, a 4- to 5-month-old puppy, and it had been buried with an elderly human. Other artifacts found at the site suggested that the location was inhabited by humans just making the transition from hunter-gathering to agriculture. Similar burial sites have been found in North America dating back over 8,000 years (Figure 2-3). Morey (2006) suggests that evaluation of the evidence associated with burial sites provides significant information about the timing of canine domestication and the special social relationship that early dogs had with humans. The early date for burial in Central North America (Illinois) suggests that dogs must have come with some of the earliest humans to settle North America, crossing the land bridge over the Bering Strait. Wherever and whenever dogs were domesticated, they quickly spread as humans moved throughout the world.

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Canine Genetics and Evolution. Modern genetic analyses have provided a new window into the domestication of the dog and its derivation from wild canids. Taxonomists had long considered the dog to be a species closely related to the various wolf subspecies (Canis lupus), but clearly separate, and were classified as Canis familiaris. In 1993, based on the resemblance of genetic, morphological, and behavioral traits, the domestic dog was reclassified as a subspecies of wolf, Canis lupus familiaris (Wozencraft, 1993). At the same time that genetic evidence can be used to verify the relation of dogs to wolves, it has been used to examine the evolutionary history of when the dog lineage separated from that of wolves.

Using genetic data to trace relationships between different species and populations, and estimating the time frame for the separation of those groups into reproductively isolated populations requires two stages. The first requires the ability to identify and sequence homologous DNA sequences found in the genomes of each population. Homologous sequences are those that are similar in form and function and are assumed to have a common evolutionary history. As two populations separate, random changes or substitutions of nucleotide bases during replication will accumulate in the genomes of each population. You can trace the lineage of a population by examining the number and order of these changes or substitutions. The fewer differences between two populations or samples, the more closely related they are. As the length of time that two populations are reproductively separated from one another increases, the number of nucleotide differences will tend to increase. If you are examining a large number of samples, this analysis rapidly grows in complexity. Genetic taxonomists have developed sophisticated statistical methods to compare substitution patterns between different samples, and determine probable relationships between the groups (for more information on molecular taxonomy see Page and Holmes, 2002).

Analysis of nucleotide substitutions can also permit estimation of time frames for the separation of two populations. It requires estimating a molecular clock. A simple description of the process requires making an estimate of how frequently nucleotide substitutions occur. If you assume that a substitution happens on average once every 5,000 years, then if two populations differ by five nucleotide substitutions in the DNA sequence being analyzed, you can estimate that the populations separated 25,000 years in the past (5,000 years multiplied by 5 substitutions). Use of this method requires a substantial number of assumptions, not the least of which is that the substitution rate has remained constant, and that your original estimate for the substitution or evolutionary rate was reasonably accurate. The DNA sequences used in an analysis may be from either nuclear-coded genes or mitochondrial DNA (mtDNA). Nuclear DNA undergoes recombination during sexual reproduction. This can introduce additional complexity into the analysis. Mitochondria are cellular organelles that contain DNA independent of the DNA found in the nucleus. This DNA will code for proteins used in the metabolic functions that occur within the mitochondria. During meiotic cell division, the mitochondria, which are found in the cytoplasm, will be shared between the resulting daughter cells. Later, during fertilization of an egg cell, the nuclear DNA of the sperm is combined with the nuclear DNA of the egg. However, the sperm do not contribute cytoplasm or mitochondria to the resulting zygote. As a result, mtDNA can be traced along matrilineal lines, from mother to daughter.

Based on an analysis of mtDNA, Vila et al. (1997) published a remarkable study adding a new twist to our thoughts on the evolution of dogs. They analyzed mtDNA sequences from 162 wolves from twenty-seven different locations around the world, 140 dogs representing 67 breeds and 5 crossbreeds, along with 5 coyotes, 2 golden jackals, 2 black-backed jackals, and 8 Simien jackals. Their resulting analysis confirmed the expectation that wolves were the ancestors of dogs. There were much greater similarities between wolf and dog mtDNA than between dog and either coyote or jackal mtDNA. There was extensive genetic diversity seen in the dogs and the authors indicate that this is likely due to domestication events from different female lineages of wolves and cross breeding between dogs and wolves multiple times during the early stages of divergence. More striking, however, was their suggestion that the wolf-dog separation may have happened as long as 135,000 years ago, which is substantially earlier than the fossil record has indicated. They postulate that the genetic divergence between dogs and wolves may have happened without substantial morphological change and that the changes seen in the fossil record are associated with a time when humans were transitioning from a hunter-gatherer existence to sedentary agrarian practices. Tsuda et al. (1997) also examined mtDNA of wolves and dogs and support the contention that dogs are descended from wolves. They also support the concept that domestication of the dog flowed from four or more female wolves, or that once domesticated, surviving dog lineages interbred multiple times with wolves.

Savolainen et al. also analyzed mtDNA sequences to evaluate the evolutionary origins of the domestic dog. They sampled 654 domestic dogs from Europe, Asia, Africa, and Arctic America along with 38 Eurasian wolves. Their results indicate that East Asian wolves were the most likely ancestors of dogs, about 15,000 years BP (2002). These data support an earlier proposition, based on fossil evidence, by Olsen and Olsen (1977), that the Chinese wolf was the ancestor of the dog. The evidence also suggests that domestication of dogs by humans in this region may have occurred multiple times.

While the archaeological evidence indicates that dogs were found in the New World by 8,000 to 9,000 years BP (Morey, 2006), the question is whether any dogs were domesticated in the New World, or were they brought over during human migration that occurred during the late Pleistocene era (Foster et al., 1996). Leonard et al. (2002) extracted mtDNA from the remains of 37 dog specimens found in the New World, deposited before the arrival of Columbus from sites in Mexico, Peru, and Bolivia. In addition, they examined remains from eleven dogs found in Alaska, but predating the arrival of European explorers in that region. They compared the nucleotide sequences they obtained to those of previously analyzed samples from wolves and dogs. Their analysis indicates that the these ancient New World dogs were not derived from New World wolves, but were in fact descended from Old World wolves. This would indicate that the humans who crossed the Bering Strait must have brought dogs with them. The level of genetic diversity observed in the ancient New World dog remains showed that these dogs must have represented a sample from a large, well-mixed, and diverse population of dogs in the Old World at that time. If the archaeological record for differentiation of the dog were accepted as being about 14,000 BP, this in turn would also suggest that dogs must have been common companions for humans as they migrated. Dogs would have moved from their origin in East Asia to the other side of the world in just a few thousand years.

While mtDNA studies have proved useful in tracing the dog's ancestry from wolves, it is less helpful for the study of different breeds of dogs. Most breeds of dogs in their current form have existed for only several hundred years (Crowley & Adelman, 1998; Wilcox & Walkowicz, 1995), and mtDNA does not evolve fast enough to generate the variation that would be required to distinguish between different breeds. Parker et al. (2004) analyzed nuclear microsatellites combined with phylogenetic analysis and genetic clustering techniques to elucidate the genetic relationships among 85 different breeds of dogs. They sampled from 414 purebred dogs and found that the variation among breeds accounts for 27% of the total genetic variation in the population studied. This is high compared to other domestic breeds and confirms the strong genetic isolation between breeds. Their cluster analysis detected four separate groupings (Figure 2-4). The first group was thought to be most ancient in origin since it also included a sample of wolves that were part of the analysis. This group may be the best living representation of the ancient canine gene pool. It included the following breeds:

* Basenji (Africa)

* Saluki and Afghan (Middle East)

* Tibetan terrier and Lhasa apso (Tibet)

* Chow chow, Pekingese, shar-pei, and shih tzu (China)

* Akita and Shiba Inu (Japan)

* Alaskan malamute, Siberian husky, and Samoyed (Arctic)

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The second group included the following:

* Mastiff, bulldog, boxer, and related breeds

* Rottweiler, Newfoundland, and Bernese mountain dog

* German shepherd dog

The third group included the following:

* Belgian sheepdog, Belgian Tervuren, Collie, Shetland sheepdog (herding)

* Irish wolfhound, greyhound, borzoi

* Saint Bernard

And a fourth group was composed of the following:

* Scent hounds

* Terriers

* Spaniels

* Pointers

* Retrievers

In many ways, the genetic clustering is consistent with our general understanding of canine breed history, and the morphological similarities observed. Wayne and Ostrander (1999) have suggested that some ancient breeds such as the dingo, New Guinea singing dog, greyhound, and mastiff may have developed in association with isolated human populations, where there was limited opportunity for them to interbreed with other types of dogs until travel become more common and easier (Figure 2-5).

An interesting result of the analysis is that several breeds generally considered among the oldest breeds of dogs, the Pharaoh hound and Ibizan hound, were not included in the first grouping of oldest breeds. While they may appear to be quite similar in appearance to the dogs depicted in Egyptian tombs, their genomes indicate that the breed must have been re-created by breeders at some more recent time by combining other breeds of dogs. Previously, genetic analysis had shown that the Xoloitzcuintli, or Mexican hairless dog, long thought to be of ancient origin, and possibly derived from New World wolves, was in fact derived from dogs originating in the Old World (Vila, Maldonado, & Wayne, 1999). Perhaps more surprising was that the "Xolo" was not related to the Chinese crested dog that it resembles physically. Continued analyses of this type will likely provide additional insights and other surprises in our understanding of dogs.

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Canine Domestication. The question is how can we reconcile the archaeological findings with the molecular genetic data and come up with a coherent picture of how wolves became dogs. A traditional view was that Man the Hunter somehow joined up with Wolf the Hunter. It was quite a romantic narrative as the two great social predators stalked large game together. Wolf puppies would sometimes be taken into caves as "pets" and over time these wolves became more and more tame. Coppinger and Coppinger provide the following simplified version of this process:

* Capture a wolf

* Tame the wolf

* Train the wolf

* Breed the wolf to other tame, trained wolves

* And, presto! A domesticated dog (2001, p. 57)

The trouble with the story is that it did not make much sense. Well-educated experts, working with extensive facilities, have their hands full trying to tame wolf puppies born in captivity (Klinghammer & Goodman, 1987). The puppies must be taken from the litter before their eyes are opened, and then bottle-fed by humans. Even then, when they have matured, they are simply more tolerant of people. They are in no way "dogs." When stressed or challenged, they will revert to their instinctive response patterns with potentially devastating consequences for a human within reach. These patterns were only papered over with a thin veneer of tameness. It is difficult to conceive that our human ancestors, at a time when they were just starting to figure out projectile weapons, would have the wherewithal to capture not just one wolf puppy and raise it to sexual maturity, but many. Selection for tameness would require a number of individuals to choose from for breeding stock. They would then pair the ones they wanted to mate, take the puppies from that litter, and several others, and do it again (and again, and again). It is more likely that any wild wolf puppies that were caught stayed around until they could escape, or the first time one snarled and snapped and then ended up as that evening's dinner entree.

The alternative process proposed by Coppinger and Coppinger (2001) goes as follows:

* People create a new niche, the village.

* Some wolves invade the new niche and gain access to a new food source.

* Those wolves that can use the new niche are genetically predisposed to show less "flight distance" than those that cannot.

* Those "tamer" wolves gain selective advantage in the new niche over the wilder ones.

This model is consistent with the archaeological evidence placing domestication of the dog at the about the same time that humans were transitioning to an agrarian lifestyle, with established villages. Villages would bring with them that unique human contribution to the environment--the dump. Leftover food, spoiled food, the bones, and the offal of animals killed and eaten would probably be tossed into a convenient location. Some wolves would come to feed at these locations. Wild wolves would flee every time someone approached the dump. Some would run a shorter distance than others; some would wait longer before they ran. These wolves would have first access to any new donations to the dump. All wild animals must acquire enough resources to reach sexual maturity, mate, and leave offspring if they are to be successful. Individuals able to exploit a resource more efficiently than others are at a distinct advantage. Those wolves most efficient at exploiting the food in the dump would not need to risk injury hunting. Over time, the flight distances for these wolves would become shorter and shorter, as individuals within this population become more and more tame. A significant part of domestication in this scenario would have taken place through natural selection.

An experiment that tested this basic scenario was carried out in Russia over a 40-year period with silver foxes, Vulpes vulpes (Trut, 1999). Dmitry Belyaev and colleagues began their work in 1959 by defining very clear selection criteria for the foxes that they would breed in succeeding generations. They started with 30 male foxes and 100 vixens. Since these foxes had already been bred in captivity for the fur trade they were already more tame than truly wild foxes, as they are adapted to being caged and fed by humans. When the first group of fox pups reached sexual maturity at seven to eight months of age they were tested for tameness. Class III foxes would flee at the approach of the experimenter, or attempt to bite when stroked or fed. Class II foxes let themselves be petted or handled, but did not show any friendly overtures to the experimenters. Class I foxes wagged their tails when petted or fed and whined for attention. The top 4 to 5 percent of male foxes and top 20 percent of vixens were chosen for breeding. By the time they reached the sixth selected generation the experimenters needed to add another class, IE, or "domesticated elite." These foxes were eager to solicit the attention of humans, and would run to greet them. While just 18 percent of the foxes were IE in the tenth generation, this had just about doubled to 35 percent in the twentieth generation, and by the time the research was reported in 1999, 70-80 percent of the population was classified as elite. These results are not surprising. There are numerous examples of successful selection for a wide variety of behaviors in many different species. In addition to the response to selection for behavioral tameness, they also observed a suite of correlated morphological changes. The foxes began to show piebald color patterns, floppy ears, and carried their tails curled up as adults, not down the way wild canid adults would normally carry their tails. They have also measured reduced adrenal cortex function, and changes in the shape of the skull. Finally, after 40 years, they have also seen some small changes in reproductive behavior. Wild foxes are strict seasonal breeders, reaching sexual maturity at about eight months of age. The selected vixens are reaching sexual maturity at about one month earlier, and are having litters that average about one pup larger. A few have even started to cycle twice per year.

What is remarkable about all of these physical changes is that these are the same general changes one would have predicted as part of a domestication scenario. However, the experimenters did not select for any physical traits, only for the operationally defined trait of tameness. If we go back to the wolves at the village garbage dump, would we see a similar set of correlated responses to "natural selection" for tameness among the wolf population feeding there? Some of the physical markers such as the tail curled up and floppy ears, would have given the humans a handy way to identify those wolves most likely to be friendly. It is a question that will deserve additional consideration in the years to come.

Once humans were able to approach and handle a domesticated wolf, it would have been possible to evaluate desirable characters, and begin selective breeding for a particular look or function. Fondon and Garner (2004) postulate that the canine genome is uniquely developed to provide high rates of genetic variation through gene-associated tandem repeat expansions and contractions, resulting in a major source of phenotypic variation. They examined a number of genes associated with developmental regulation and found that in most of the cases, allelic variations were a function of incremental differences in length, generally corresponding to two or three repeats of the length of the most common allele for that locus. These sorts of alterations at genes associated with the control of development events could have wide-ranging effects on the morphological development of dogs and could have accounted for the rapid evolution of highly diverse breeds shortly after the domestication of the dog from the wolf.

Another interesting view of dog evolution by McGhee (2002) brings us back full circle to the initial premise of Caras (1996) that the domestication process was a form of co-evolution of animals and humans. McGhee suggests that if the oldest dates of over 100,000 BP for separation of wolves and dogs were true, then those early "dogs" would have lived in concert with humans just barely more advanced socially than they were. These early human groups, only recently evolved from Homo erectus, probably lived in small, mobile family bands, organized in a way similar to modern chimpanzee bands. These wolf and human family groups may have coalesced among many of the same required resources, water, and food. These mobile bands of humans would not have left food or debris in any quantity in a single location as they hunted and gathered what they could. While these wolf-dogs may have accumulated the genetic signatures we can detect with modern analytic methods, they likely did not manifest anatomical changes that could be reflected in the fossil record. Once small groups of humans began to gather and stay in a particular location for extended periods of time, beginning the transition from hunter-gatherer to farmer-villager, this population of wolves would have been pre-adapted to begin the rapid changes in behavior and morphology that would allow them to exploit this new human-generated niche. McGhee raises the question of whether, in addition to the influence humans obviously had on the evolution of some wolves to become dogs, wolves influenced our own evolution. Did the human hunter bands that might be associated with wolf packs enjoy an advantage? Were these the groups that were most likely to thrive and go on to form the first villages, where wolves truly became dogs? He suggests that some elements of this story may be recorded in human folklore. There are numerous stories of wolves nurturing and caring for humans. Romulus and Remus, the founders of Rome were supposedly fostered by a she-wolf. He also sees that our frequent vilification and persecution of wolves by humans is part of this same close relationship. Our cultural depictions of the wolf are seldom neutral, being either villain or savior, and may be rooted in a history between humans and wolves that predates history and culture.

Social History of Dogs. If we accept that dogs were both genetically and physically differentiated from wolves by 12,000-14,000 year ago, it would be another 10,000 years until the next major steps in the development in dogs took place. Until this time dogs were pretty much a generic lot, probably very similar in size and appearance to the pariah dogs still found around the world today. In addition to scavenging around human villages they probably went along on hunting forays, helping to raise game, chase, and either bring down or locate wounded animals. They also would have proved useful helping to guard the next animals domesticated by humans, sheep and goats. Their natural territoriality and social and protective behaviors would have lent themselves to these tasks. In was not until 5,000-6,000 years ago we see the first evidence for development of dogs for specific purposes (Rogers, 2005). The simple village of humans has become far more sophisticated and different people might have different roles or tasks. Hierarchies and social structures had developed to the point that there was now a ruling class. This upper class no longer needed to raise or catch their own food. They depended on others for the basic needs. Hunting now became a form of recreation and the implements of the hunt were a sign of wealth and power. Among these signs were dogs, specially bred and raised for the hunt. The first of these were dogs very much like the salukis of today. Evidence of these dogs is found in the art and other artifacts of ancient Sumeria and Mesopotamia. These first breeds were long and lean with deep chests. They were sight hounds and beautifully adapted for chasing game on the open deserts. The hunters, mostly nobility, would ride out to the hunt in chariots with dogs kept on leashes until game was spotted. Once small antelope or other game was spotted the dogs would be let loose to chase and bring down the quarry.

The next type of dog to appear was the large, heavy-boned mastiff. There is evidence that mastiffs may have appeared independently in both northern India or Tibet and somewhere in the Mediterranean. Five thousand years ago the Island of Molossis, near Greece, was known for such dogs and for some time these large dogs were called Molossians. They were used as guards, hunting large dangerous game such as lions, and also for warfare. These war dogs might be outfitted with armor, and have a collar of blades. As they ran slashing an enemy's lines they would wound and cripple men and horses. At other times they were outfitted with torches and the flames would frighten horses and start brush and other materials on fire. By 4,000 years ago, the Egyptians had their own variety of sight hound, very similar to present day greyhounds. They also had mastiffs and here we now see the first small dogs. The Egyptians also memorialized their dogs in their art, and as part of their religion (Wilcox & Walkowicz, 1995). Anubis, the Egyptian god of the dead, is depicted with the body of a man and the head of a dog. It was Anubis who would gather the dead into his arms and transport them to the next world. The Egyptians were highly dependent on the annual flooding of the Nile River for continual renewal of their fertile croplands, the star that would presage this event each year was named Sirius, the Dog Star. The appearance of Sirius would forewarn the shepherds to move their flocks to higher ground.

The next type of dog to develop was the scent hound. These hounds were derived from the mastiffs. Some of the lighter, quicker forms were selected to trail game by scent. The loose skin and floppy ears of hounds reflect this heritage. The Greeks had highly developed hounds for hunting which they called Laconians. The Greeks respected the loyalty and honesty of dogs and Homer highlights this as part of the climax of his epic Odyssey (Rogers, 2005). When Odysseus finally makes it back to Ithaca after the exceptional ordeal that he and his men endured in the return from Troy, he finds that his home has been taken over by interlopers. Bedraggled and disheveled, he is not recognized by the members of his household. It is Argus, his loyal hound now old and discarded on a dung heap, that recognizes his master. Barely able to move, Argus shows his happiness in the return of Odysseus silently, before passing away. Odysseus, who would soon slaughter the interlopers who sought his kingdom, wealth, and hand of his wife, stifled a tear in gratitude for such love and loyalty. Unlike the Egyptians, the Greeks dreaded the dog days of summer signaled by the rise of Sirius. For them, these days were times of fever, sunstroke, and drought. They believed that dogs went mad, and a dog bite at this time would result in rabies.

The Romans, of course, traced the foundation of their culture to the twins Romulus and Remus, who were suckled by a she-wolf. They began to divide dogs into various groups depending on their use. They recognized the following common categories for dogs:

* House

* Shepherd

* Sporting

* Pugnacious or war

* Dogs that ran by scent

* Dogs that ran on sight (Wilcox & Walkowicz, 1995).

The Romans were particularly fond of their war dogs and used them not only in combat, but in the Coliseum as well. These dogs would battle one another, men, or wild animals such as lions and bears. When Julius Caesar invaded Britain, he brought along a great number of war dogs. Once there, he was impressed with the large, aggressive dogs that the local inhabitants possessed and also brought to battle. When the Romans found especially large and powerful dogs on an island in the Mediterranean Sea, they named the island after the dogs--the Canis or what became the Canary Islands. Later, the small songbirds found on the islands would be called canaries. The Romans were apparently fond of small house dogs as well, and kept miniature or toy dogs, among the first being the Maltese. The remains of many of different sizes were found in the ruins of Pompeii. Among these were the skeleton of an elderly person with their arms wrapped around the skeleton of what was likely a cherished pet.

During the Middle Ages hunting became very sophisticated as a pastime. It was closely linked to feudal lords and the ladies and knights of their courts. Many new breeds and types of dogs were developed for hunting specific types of game. Hunting might be done on horseback or afoot. Among the most popular animals to hunt were various deer and boar. Boar were a particular challenge since they might seek cover in thickets and were capable of doing significant damage to hound, horse, and huntsman with their tusks. The huntsman went afield with lance and spear, and hunting was pursued to develop the skills with arms and horseback needed for battle. Small game such as rabbits and hares might be taken by coursing with sight hounds. It was not uncommon to hunt with both dogs and falcons. The dogs would drive the game into the open where the falcon could stoop on it. Birds could also be taken in this way. By 1486, Juliana Barnes, the prioress of Sopwell Nunnery listed the following breeds in the Boke of St. Albans:

* Greyhound

* Bastard

* Mengrell

* Mastif

* Lemor

* Raches

* Kenettys

* Terroures

* Butcher's Hounds

* Dunghill Dogges

* Tyrndeltaylles

* Prycheryd Currys

* Small Ladyes Poppees That Bere Awayethe the Flees (Wilcox & Walkowicz, 1995).

These breeds included many dogs that would be familiar to us today: greyhounds, mastiffs, small and large scent hounds, terriers, bulldogs, and mongrels. The Small Ladyes Poppees were lap dogs, and in addition to companionship, and keeping someone warm in the drafty castles, were presumed to attract the fleas common to everyone to jump off the ladies and onto them. The next type of dog developed during this era was the spaniel. It was a bird specialist and would locate and then either hold the birds in place to be netted, or flushed for the falcons.

While the many hunting dogs, guard dogs, and lap dogs of the wealthy might be pampered, the dogs of the poor serfs shared their limited means and circumstances. Terriers were a particular help as they were small, aggressive dogs that needed little upkeep and were deadly efficient in ridding a field of badgers or barn of rats. A particularly onerous treatment was reserved for the dogs belonging to any peasants that lived near the protected estates and hunting lands of the lord. While it was accepted that dogs were needed to guard flocks and perform other tasks, the lords were so jealous of their hunting and game, steps were taken to ensure that peasant dogs would be unable to pursue or kill game. It was therefore not uncommon for the tendons in the hind legs to be cut, or the toes on the front feet. Thus hobbled, these dogs could limp about through their daily tasks, but were unable to threaten the royal deer. Peasants could not own hunting dogs. However, poachers developed their own dog, the lurcher. A cross between a greyhound and either a terrier or sheepdog, the lurcher was quick, silent, deadly, and efficient in its pursuit of small game that a poacher depended on. Lurchers are still bred and used for coursing hare and rabbit.

As the great era of European exploration of the world began at the end of the fifteenth century and on through the next several centuries, their dogs came with them. The Spanish brought fierce mastiffs with them to the New World and used them to intimidate the natives. On their return home, in addition to gold, spices, and other riches, they brought back some of the dogs they found on their travels. This resulted in additional new breeds being introduced to Europe, and their crosses with the existing breeds led to a proliferation of additional new breeds being developed.

The current culture of purebred dogs, the dog fancy, and the formal competitions for dogs developed during the 1800s. As the role of royalty and hereditary nobles began to decline in social influence, the growing wealth of merchants, financiers, and industrial magnates began to dominate. Among the many habits of the upper classes that they accepted and continued was an interest in dogs. Owning and breeding purebred dogs was a sign of refinement and a respected pastime for a Victorian gentleman. Informal shows, competitions, and exhibits were held to demonstrate the qualities of the top dogs. In 1859, the first formal show was held in Newcastle, England. It was sponsored by a gun maker, and was limited to bird dogs. The winning exhibitors were awarded guns as prizes by the sponsor. The Westminster Kennel Club held their first show at Madison Square Garden in New York in 1877. Henry Bergh, founder of The American Society for the Prevention of Cruelty to Animals (ASPCA), was keynote speaker (Figure 2-6). He praised the owners for their dedication to dogs and the dogs for their devotion to humans: "They aim at the improvement of the race of animals which you are invited here tonight to inspect.... He is dedicated to his master, adopts his manners, defends his property, attaches himself to him until death, and all this springs not from necessity or restraint, but simple gratitude and true friendship." Bergh also called upon those in attendance to see to the welfare of all dogs, whether pedigreed or not. Accounts of the event suggest that few people may have heard Bergh's remarks or those of the other speakers as they were overwhelmed by the din of barking and howling canines. Nonetheless, the kennel club responded to Bergh by donating proceeds from the show to the ASPCA to provide care for stray dogs and those belonging to the poor and indigent (ASPCA Archives, 1877 press clips).

[FIGURE 2-6 OMITTED]

Much like the feudal lords who preceded them in the care of dogs, this developing class of dog fanciers could and would pamper their dogs. However, as in times past, dogs still suffered the lot of their masters. While dogs belonging to the wealthy might have special beds, and choice meals, dogs belonging to the poor needed to make do with what they might scavenge and scrounge. Instead of living in villages on the lands of a feudal lord, the poor were now often found crowded into cities. Their dogs were not needed to help with herding or guarding as many of these people worked in factories or manufacturing establishments. Others might be small vendors selling vegetables, odds and ends, or collecting rags. Those too poor to afford a horse to pull their carts would harness a dog to their carts. In many cases at the end of the day, the dog would be let loose to find his own food by scavenging through the streets and gutters. The wretched condition of these dogs was one concern of the early humane groups working to prevent cruelty to animals at this time (ASPCA Archives, Buffet notes; also see Chapter 2.4). There were small dogs called turnspits that trotted within wheels that turned the gears of rotisseries in shops and restaurants. Other dogs were used in the fighting pit against other dogs, wild animals, or rats. Still uncounted numbers wandered the streets where they harassed people and horses, and suffered. Every so often, when rabies might appear in a city's stray dogs, hundreds or thousands might be rounded up and killed by drowning, clubbing, or starvation.

During this same era dogs would be glorified for many of the same traits that endear them to people today: loyalty, courage, and honesty. One of the most famous stories from the early part of this era is that of Greyfriars Bobby. Bobby's master was a market policeman in Edinburgh. When his master died and was buried at Greyfriars Churchyard, Bobby did not go on to live with another master. Instead, for fourteen years he lived around the churchyard, surviving on the handouts of shopkeepers and townsfolk. Journalists and authors of the age wrote heartwarming stories about Bobby's love and loyalty to his master. He was eventually honored with a bronze statue in 1873 (Rogers, 2005). Loyalty has remained a popular theme in literature about dogs, and in 2004, two stories, one nonfiction and one fiction, were published for children about Hachiko (Newman, 2004; Turner, 2004). This dog belonged to a Japanese college professor who took a train to work each day. His dog, Hachiko, would follow him to the train station in the morning and then return home. In the afternoon, he returned to the station to wait for the professor. The professor died suddenly at work one day, and Hachiko loyally returned to the station each day to await the professor's return. This went on for almost ten years until Hachiko died. He is now remembered with a statue in the Shibuya train station.

The kennel clubs that formed sought to highlight the refined nature of dogs, those that were the very best representatives of their type. Purebred dogs were typically the passion of individuals who kept their own packs of hounds, or bird dogs, for hunting or some other type of dog that held a particular interest for them. They would breed and maintain their own records or studbooks. These records might be shared with other enthusiasts interested in the same type of dogs. Most of all, they bred for performance, doing the job that the breed was originally developed to perform. Each breeder would strive for some uniformity in size and appearance. This was particularly important in dogs that would work in packs, where the efficiency and aesthetics of the pack were dependent upon uniform size and speed, and even matching the color patterns and harmonizing the voices in the pack as it trailed game (Black, 1949). These breeders would outcross or breed their dogs to other lines or varieties of the same breed, or sometimes to other related breeds to add particular qualities or to correct faults. The formation of the American Kennel Club (AKC) in 1884 would bring written standards to the dog fancy. These standards would be developed and agreed upon by individual breed clubs, and these clubs together would form the AKC. It was no longer a single breeder looking to establish uniformity in the dogs that he or she bred. The goal would be to ensure that all registered dogs of that breed would have a uniform quality and appearance based on the written standards. Judging dogs at shows or field competitions would be based on the same standard. The development of these formal rules also resulted in the closure of the studbooks. Registration of a dog would now require that parents be registered.

In the years after World War II, a burgeoning middle class moved into the new suburbs outside the cities. There, with room and extra income, they completed their families by getting a dog. Walt Disney films like Lady and the Tramp and Old Yeller made it clear that dogs were a desirable and important part of families. These families now had televisions, and the post-war years brought them stories of Lassie, Rin Tin Tin, and the family fare of Walt Disney's weekly television shows, many of which featured dogs. In a marketing culture that now celebrated name brand products, purebred dogs became the name brand for dogs. The small group of fanciers and breeders working with the various breeds were unable to meet the new demand, and commercial breeding establishments were created to meet the need for pet dogs for retail sale. These puppy mills eventually became a significant concern for animal welfare groups, and there have been a number of campaigns conducted to regulate or eliminate them (see Controversies).

Feline Domestication

One story of the cat's origin has them coming forth from the sneeze of a lion on Noah's ark (Simpson, 1903). This explanation highlights two interesting aspects of the cat: that it was a late arrival in the family of animals (all of the other animals were already on the ark) and the remarkable similarity in behavior that the cat retains with its wild relatives. The domestication of cats is a more recent historical event than that of dogs. While there is some evidence that tamed wildcats may have been kept as "pets" as long as 9,500 years ago in Cyprus (Vigne et al., 2004) and 6,000 years ago elsewhere (Davis, 1987), their development as a domesticated species seems to be pretty well established in Egypt about 4,000 years ago (Mery, 1968). There has been some debate about the ancestral species that gave rise to the domestic cat, Felis catus. It has been difficult to track the evolution of the domestic cat since there is such a high degree of similarity in the morphology among the various small wildcat species, and there has been limited morphological differentiation between domestic cats and the most likely ancestral species. Several different lines of thought all converge on the African wildcat, Felis sylvestris lybica, as the most likely ancestor.

Cameron-Beaumont, Lowe, and Bradshaw (2002) performed a comparative behavioral survey of sixteen species and subspecies of small wildcats kept in zoos. They arranged for zookeepers to complete a survey form on the tameness and affiliative behaviors of cats in their care. They hoped to identify wild species that might have had a higher frequency of behaviors that would have lent themselves to the domestication process. They collected data on whether keepers were able to enter the cats' cages, and the extent to which the cats showed friendly behaviors similar to those showed by domestic cats such as rubbing or licking the keeper. Their results showed that the expression of affiliative behavior toward people was patchy, and widely spread among the species evaluated. This suggests that several of them could have been candidates for domestication. However, they conclude, "the tendency toward tameness among the small felids points for a localized human need as being the primary reason for the domestication of F. s. lybica, rather than any special features of its behavioral biology" (p. 365). Serpell (2000) indicates that the European wildcats, Felis sylvestris sylvestris, have a reputation for fierceness and are difficult to tame, even when reared as kittens. This is borne out of the data presented by Cameron-Beaumont et al.

Randi and Ragni (1991) analyzed populations of F. sylvestris, lybica, and catus populations from Sardinia, Sicily, and the Italian mainland and based on comparisons of morphometrics and allozyme variation concluded that lybica was the most likely ancestor of the domestic cat. More recently, Johnson et al. (2006) evaluated molecular genetic data related to cat evolution. Modern felid lineages appeared 11 million years ago in Asia. There was a substantial amount of movement, with at least ten intercontinental migrations associated with historic changes in sea level, allowing ancestral cat populations to move from one continent to another over land bridges. Cats were very successful predators, second only to humans. The lineage leading to the domestic cat appeared in Africa and Asia about 6.2 million years ago (Figure 2-7). It may have arisen from a family that had migrated to North America and then returned. A further branch appears in this lineage about 1 million years ago, leading to a family of small cats, and eventually to the ancestors of the domestic cat.

A further analysis of both domestic cat DNA samples (mtDNA), along with samples from five different subspecies of Felis sylvestris confirmed that cats were domesticated in the Near East (Driscoll et al., 2007). This likely happened in the Fertile Crescent region during the time when agricultural villages were developing about 9,000 years BP. There were at least five different founding females from across that region, and the resulting lineages were subsequently transported worldwide with human assistance.

In many ways, the archaeological evidence associating cat remains with human remains in ancient Egypt may be the most convincing evidence that F. sylvestris lybica gave rise to the modern domestic cat. A small tomb dated to about 2000 BCE yielded the bones of seventeen cats when it was excavated (Malek, 1993; Mery, 1968). Egypt was home to a rapidly expanding agricultural economy. Each year the Nile would flood and then recede to leave a rich layer of fertile silt to renew the fields. Grain was harvested in great quantities and stored against times of need. These stores attracted rats and mice, the natural prey of the wild cat, F. s. lybica. It is likely that the cat first came to live near people to prey upon the rodents. Over time, it is likely that they adapted to living in closer and closer proximity to humans. Ancient Egyptian religious practices assigned roles to a wide variety of animals and it would only be natural for cats that now protected their grain to be absorbed into these representations (Malek, 1993; Mellen, 1940). Mellen notes that the Egyptians had a history of taming a variety of creatures, including hawks, ibis, and others, and the cat would also attract their efforts.

[FIGURE 2-7 OMITTED]

Social Evolution of the Cat. One manifestation of the Egyptian sun god Ra was as a male cat. Each night, in this form, Ra would battle the serpent of darkness (Malek, 1993). The Egyptian Book of the Dead mentions cats, and coffins from the Sixth Dynasty (2600 BCE) were inscribed with language from the book and painted with cat figures (Mellen, 1940). Eventually the cat would transition from primary representation as a male figure to an association with females. This would culminate during the Twenty-second Dynasty, 945-715 BCE (Serpell, 2000; p. 184), with a rise in the prominence of the goddess Bastet (Figure 2-8). Bastet had long been associated with the city of Bubastis. Egypt was a nation of clans and tribes, each with their own totemic animal (Mellen, 1940; Malek, 1993) and Bubastis was a center of cat worship, and their totem was Bastet (or Pasht). When a family from Bubastis came to prominence in ruling Egypt, they also brought their totem to prominence with them. Bastet was associated with fertility and plenty and the annual festival in her honor was popular and celebrated with vigor (Mellen, 1940). Serpell (2000) suggests that during this era, the cat held a place in Egyptian society equivalent to that held by the cow in present-day India. Many families owned cats and the death of the family cat would throw the family into mourning, shaving their eyebrows in a traditional sign of respect. Families that could afford the price would arrange to have their cat embalmed and buried in special cat cemeteries. Killing a cat was prohibited and might be punished by death. Mobs could become so enraged by the killing of a cat that they would fall upon the perpetrator and tear them to pieces (Mellen, 1940). In a strange twist, temple catteries were maintained for the purpose of providing cats for sacrifice and votive offerings (Armitage & Clutton-Brock, 1981). It was a great honor to care for these cats, and several species of catfish were raised in special ponds to feed these sacred cats (Mellen, 1940). However, before they reached the age of two they were strangled or killed by breaking their necks and then embalmed to be offered as votives during worship.

[FIGURE 2-8 OMITTED]

The Egyptians prohibited the export of cats (Mery, 1968), but Phoenician traders eventually carried cats to other areas where they traded (Mellen, 1940). By about 500 BCE, a cat appears on a marble block in Greece (Zeuner, 1963). Greeks were responsible for transporting cats to Rome. Neither the Greeks nor the Romans took the cat to heart as they did dogs. Mery (1968, p. 22) points out that while dog bones were found at Pompeii, no cat bones were recovered. Regardless, the Romans were likely responsible for introducing cats to the rest of Europe, Britain, and throughout their Empire (Zeuner, 1963). The cat appears again in a place of honor among the Norse in their mythology. Freyja, the goddess of love, like Bastet, was drawn in a cart by two cats. Norse maidens married if possible on Freyja's day (Friday), and if the sun shone during the ceremony it was a good omen and guests would note that the bride had "taken good care of cat" (Bulfinch, 1979; Mellen, 1940, p. 224).

The rise of Christianity in Europe during the Middle Ages brought hard times for cats. Extensive efforts were made to stamp out heretical and unorthodox teachings. Between the twelfth and fourteenth centuries, cat and devil worship were linked in attacks on various sects and individuals (Serpell, 2000). The lithe, sensuous behavior of the cat that lent itself to identification with female goddesses of fertility now brought the ire of a conservative, male, clerical hierarchy intent on erasing links to a past that were not consistent with their vision of orthodoxy (Russell, 1972). The next several centuries saw cats linked to the practice of witchcraft. Witches were variously accused of flying to their gatherings on the backs of demons in the form of black cats, using cats as their familiars or demonic companions and even taking on the shape of black cats themselves (Mellen, 1940; Mery, 1968; Russell, 1972). These many negative associations opened cats to a wide range of mistreatment and persecution throughout Europe in the Middle Ages (Lockwood, 2005). Cats were burned, strangled, and beaten to death in a wide range of traditional events and activities meant to "kill" or drive off the devil. This was in addition to the many casual cruelties visited upon the uncounted cats that roamed the streets of European cities. An interesting contradiction to this linkage of black cats and evil were the matagots in several parts of France. These were black cats that brought riches if they were brought into the house, loved, and fed (Mery, 1968).

Dislike and fear of cats was not restricted to the clerics and poorly educated peasants. Georges Buffon, the premier naturalist of the eighteenth and nineteenth centuries had a great appreciation for dogs (Rogers, 2005). Buffon was especially enamored with a dog's submissive nature, loyalty, and willingness to please its master. Apparently the independent nature of cats, combined with their "insatiable" sexuality, was not to Buffon's liking, and he went out of his way to revile them (Kete, 1994; Simpson, 1903).

While witches and black cats remain linked in the jovial celebration of Halloween in modern times, a whiff of superstition and concern remains, but this time to protect cats. Each year, at the end of October, it is not unusual for animal shelters to "hide" any black cats in their facilities until after Halloween, protecting them from adoption by people with evil intent. This happens in the absence of reasonable data to support this particular concern.

Development of Cat Fancy. In spite of the many challenges that confronted cats and the people who liked them, by the end of the nineteenth century an active cat fancy was in development. The first cat show was organized by Harrison Weir, and held at the Crystal Palace in London in 1871 (Simpson, 1903). Harrison went on to help organize and become president of the National Cat Club (NCC) in 1887. The NCC was organized for the following:

1. Promote honesty in the breeding of cats, in each breed and variety.

2. Determine the classification required for each breed or variety, and encourage the adoption of such classifications by breeders, exhibitors, judges, and the committees of all cat shows.

3. Maintain and keep the National Register of Cats.

4. Assist the showing and breeding of cats by holding cat shows under the best sanitary conditions, giving Championships and other prizes, and otherwise in all its power to protect and advance the interest of cats and their owners.

At the time, it cost 1 shilling to register a cat with the National Cat Club, and 5 shillings to enter it into the studbook (Simpson, 1903).

The first major American cat show was held in Madison Square Garden on May 8, 1895. It was organized by an Englishman, James T. Hyde, who had previously arranged horse shows at the Garden. He had visited the Crystal Palace cat show in England and imported the concept to the United States. This first show was won by a Maine coon cat named Cosie, owned by Mrs. Fred Brown (Figure 2-9). Among the prizes were a silver collar and a medal (Thomas, 2004). It was certainly fitting to have a Maine coon cat win that first American show. Maine coon cats were a breed developed in the United States and already had their own registry and competitions in Maine (Miller, 2004). A variety of cat clubs began to crop up in the United States during the years around the turn of the century. The first of these was the Beresford Cat Club of Chicago in 1899 (Thomas, 2004). The American Cat Association (ACA) formed in 1901 to provide a registry and pedigree service. There was a split in 1906 because of differences over registration policies, and the Cat Fanciers' Association was formed. The current form of the CFA came about in 1955 as an association of cat clubs.

The Early Cat Fancy. Simpson's book is a comprehensive look at the world of the cat fancy at the turn of the century (Simpson, 1903). She presents a variety of opinions and practices that remain common to this day. She begins by stating that "dogs are more essentially the friends of men, and cats may be considered as the chosen allies of womankind" (p. vii). This remains a common attitude and tends to be reflected in different ways that males and females interact with cats (Turner, 2000). Simpson goes on to point out that there was already a remarkable array of "catty" Christmas souvenirs, almanacs, calendars, and various other objects that would appeal to the cat lover. There was also a wide range of written materials for those involved with breeding and showing cats. Fur and Feather included information on cats, birds, rabbits, poultry, cavies, and mice, indicating that not just cats were popular as pets. Our Cats promoted itself as "the only newspaper in the world solely devoted to cats." In the United States, the Cat Journal, the Pet Stock News, and Field and Fancy all provided support for the growing interest in cats. The Cat Courier, except for some of the prices and trade names, is quite similar to the periodicals available to cat fanciers' today (Figure 2-10). It included ads for patent medicine for cats, Spratt's cat foods, the schedule for upcoming shows and the results of recent shows, and obituaries for well-known cat fanciers and highly successful and beloved cats. Studs were advertised at $10-$15, payable at the time of service, and pedigreed kittens could be purchased for $10-$25.

[FIGURE 2-9 OMITTED]

Simpson also provides a rare glimpse into the details of caring for pedigreed cats at the time (Simpson, 1903). This is of particular interest since commercial foods were not generally available, and veterinary care for cats was limited. She states that while a mix of table scraps would be perfectly fine for a couple of pet cats, caring for a number of cats in a cattery required some attention to a scheduled diet. She then gave an example of the diet that she followed for her own cattery:

* 2 days a week--fish with some rice or Freeman's Scientific Diet

* 2-3 days a week--raw meat

* 2 days a week--horsemeat

* 1 day--Spratt's biscuit soaked with stock

* 1 day--vegetables to purify the blood and keep the bowels in good condition. (pp. 37-57)

[FIGURE 2-10 OMITTED]

The cattery should have several four-inch deep metal tins filled with earth for the cats to use for elimination. Sand, ashes, paper, and sawdust were other common fillers used in cat boxes until the invention of cat litter in the 1950s.

She recommends that male cats that will not be used for shows and breeding be neutered at five to eight months of age, since keeping a male in the house after ten months of age would be unpleasant due to spraying to mark territories. Females should be neutered at a bit older age. Simpson points out that these altered cats will make great pets, and that they should receive greater attention at the various cat shows. Most interesting is that she also recommends that the active breeder install their own humane euthanasia box for humane killing of sick, old, or injured cats and kittens.

Cat breeds at this time were usually regional varieties with a unique appearance and known pedigree or history. As various cat populations were discovered in different parts of the world, they would eventually make their way to a serious cat breeder who would take on the task of standardizing the appearance of the "breed" and introducing it to other cat fanciers. The first documented, deliberate attempt to create a new breed did not happen until the 1920s, when the Himalayan was created by crossing stocky, longhaired Persian cats with the shorthaired, slender Siamese. After a number of generations, the popular Himalayan was accepted as a new breed. There are now three generally accepted ways for a new breed to be created:

1. Selective breeding to enhance a national or regional variety (these are often called natural breeds).

2. Hybridization of two or more different breeds. The Himalayan would be an example of this. There have been several breeds recently developed by crossing domestic cats with various small wild cats.

3. Mutation. The Scottish fold and munchkin cats are the result of isolated, spontaneous mutations that resulted in folded ears and shortened legs respectively that were preserved through breeding. (Thomas, 2004, pp. 3-13)

The Modern Cat Fancy. The Cat Fanciers' Association is a leading organization in the world of modern cat breeding, showing, and care. Its first two shows were held in 1906 in Detroit and Buffalo, and it now sanctions nearly 400 shows each year. The first cat registered by the Cat Fanciers' Association (CFA) was an orange and white longhaired male named Peter, born on May 2, 1906. By 1987 the CFA registered its billionth cat, Grand Champion Tosa Mahogany (Thomas, 2004). The CFA had undergone a variety of changes to reflect the needs of its constituency and currently adheres to the following objectives:

* The promotion of the welfare of cats and the improvement of their breed

* The registration of pedigrees of cats and kittens

* The promulgation of rules for the management of cat shows

* The licensing of cats shows held under the rules of this organization

* The promotion of interests of breeders and exhibitors of cats (Cat Fanciers' Association, 2003, p. 2)

Horses

The ancestors of what is the modern horse (Equus caballus) began its evolution 54 million years ago in North America. These primitive relatives were distinctly unlike the large, sleek animals that capture the affection and admiration of so many people and cultures of today. The Eohippus was about the size of a fox and had four toes on its front feet and three on its hind feet. By 35 million BP, Mesohippus was a larger animal and it now had three toes on the front feet, with an enlarged middle toe. As the forests of that era began to thin, and with more grasslands, the Mesohippus retained the teeth of a browser. As the grasslands became a more dominant part of the landscape, the Merychippus of 20 million BP had grown to 35 inches in height and its teeth were now adapted for grazing. The enlarged middle toe was now a weight-bearing hoof. The quick, social horse ancestor would eventually become a true monodactyl, or one-toed animal. That single toe had become a true hoof, providing excellent speed to evade predators on the wide-open grasslands. Pliohippus was highly successful and quickly multiplied and spread across a wide region. Some groups even crossed fromNorth America into Asia over the Bering land bridge. This would prove significant for survival of the evolutionary line. As climates changed again, the original equines disappeared from the Western Hemisphere.

The ancestors that crossed into Asia rapidly spread through that continent and into Europe. They made quite an impression upon the early humans that lived at that time. Horses are among the most commonly depicted animals in early cave paintings. It was clear that humans valued horses as a food source, and the paintings frequently depicted hunting scenes with horses as the quarry. The presence of butchered bones in caves and at the base of cliffs, where herds may have been driven as part of the hunting strategy, provide strong ample evidence of this appreciation. When domestication of the horse began, probably in the region of the Asian steppes north of the Black Sea, it was as a source of food rather than transportation (Clutton-Brock, 1981).

Oxen were already being used for draft purposes in the Near East. As horses were introduced, the harnesses and yokes used with oxen were adapted for use with them. It took some time before horses found their true place in partnership with humans. Until wheeled carts and wagons were readily available, the stronger, slower oxen were better suited to hauling the sledges used to transport goods and materials. The rapid ascent of horses in value and appreciation came when they were put in front of light, wheeled, war chariots. Armies that employed horses in this way were dominant on the battlefield, and for centuries to come, horses were a fundamental element of every successful military force. Their value was so great that ownership was often limited to royalty and ruling classes. It was not until about 3,000 BP that intrepid souls made riding horses a common practice. By this time, kings and conquerors searched their lands and beyond for high-quality horses. The specialization of horses also began. Swift, spirited horses were selected and bred for war, and large, placid horses were bred for draft purposes. In nearly every place where horses were taken, they adapted to the conditions and needs of the people.

In 1519, the horse had its homecoming. Coronado brought 150 horses with him when he sailed to the New World to explore, conquer, and search for gold and other treasures. The natives they encountered were terrified of the massive, snorting beasts that Coronado and his men rode. Knowing their value, Coronado and the other conquistadors that followed prohibited the Indians from possessing horses. It was not long, however, before some horses were lost and others were stolen or otherwise ended up on their own. These horses found the land of their ancestors to their liking and once again wandered the grasslands of the Western Hemisphere. Along with horses that would come from settlers moving across the continent, strays from herds being moved, and those lost by military units, a new population of wild horses would again roam North America. Often called mustangs, these are not true wild horses, as they are descended from domestic stock. The only remaining true wild horses in the world are the Przewalski horses of Mongolia (Equus przewalski). In appearance, the Przewalski horse resembles the horses that appear in the ancient cave drawings. Short and stocky with brushy manes, they remain a link to an earlier era before horses were domesticated. These horses are now rare, and many zoos around the world have worked to breed them in captivity to preserve the species (Parker, 2003).

The role of the horse in modern times continues to evolve. First seen as a source of food, the horse eventually became the primary form of land transportation for humans from the time of the Pharaohs until the early part of the twentieth century. An aura developed around the breeding and care of horses unlike that for any other domestic animal. They were pampered and praised. Great racehorses were celebrities, as well known (or better) than any human athletes of their time. At the same time, their working brethren were beaten and driven until they died in their labors. It was no coincidence that it was the treatment of horses that gave rise to the humane movement in the United States. The role and place of horses in our modern society continues to evolve. While many horses still earn a living working on ranches and in other occupations, others are accepting the mantle of companions. They are fed and sheltered for the pleasure of their human companions. They now earn their living not by hauling loads, transporting people, racing, or performing. They are horses to ride for pleasure, not for work. This transition of horses from the role of livestock and work animals to companions is still evolving, and certainly not universal in our society. Debates on the use of horses for pulling carriages in the city, in rodeos, and other activities often generate intense confrontations. Perhaps nothing signifies the crux of this evolution as the debate over the slaughter of unwanted horses for consumption (see hot button issues). For people who see horses as an investment for work, racing, or other function, selling a horse for slaughter at the end of its useful life is a reasonable way to recoup some additional value. They will argue that horses have been on the menu for the entire known history of the association between humans and horses, which in fact predates other uses and functions for the horse. For others however, this is clearly beyond the pale. The horse is more than just another animal that we breed and use and extract final value.

Small Animals

Often called shelf pets or pocket pets, a variety of small animals have become common and popular in the home. While it is likely that people have been catching and keeping a variety of small animals as pets for a large part of human history, the domestication and adaptation of small animals as companions is a more recent development. In some cases these animals were originally used for another function. They might have been used as food at one time, or as subjects for laboratory research. What they generally share is small size, shorter life spans than dogs or cats, and less intense care requirements. Unfortunately, it is sometimes the case that people overlook the special needs that many of these species require. They may require very specific diets and environments.

Ferrets. Ferrets currently kept as pets have descended from some form of polecat (Musleta sp.). More than likely it was the European polecat (Mustela putorius; Hemmer, 1990). The Romans used ferrets and probably domesticated them in the Mediterranean region about 3,000 years BP (Clutton-Brock, 1999). They were originally used to hunt rabbits and rats. Their slender bodies and quickness allowed them to drive rabbits out their burrows (Figure 2-11). Their skill and usefulness for this purpose are memorialized in the use of the phrase "ferret out" to describe digging into some topic to find and answer. Ferrets were brought to the New World in the seventeenth century to control rats. They did not become popular as pets until the 1970s. Ferrets are now bred in several different color forms. They are generally fastidious in their grooming, and can be litter-trained. Their size and cleanliness make them a desirable apartment pet. They are not welcome everywhere, however. New York City, the state of California, and other locales prohibit possession of ferrets. This apparently arises from concerns about their potential danger to children--they will bite sometimes--and their possible release and establishment as an introduced species that could threaten native birds and animals.

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Rabbits. Rabbits (Oryctolagus cuniculus) originated in Europe and were probably domesticated during the Middle Ages (Gendron, 2000). The Romans, however, may have penned wild rabbits to fatten them before slaughter. Monks played an important role in breeding and domesticating rabbits. It is thought that one reason for the growth in the popularity in raising rabbits during the Middle Ages is that church teachings held that the unborn baby rabbits were not considered meat and could be eaten on days of abstinence when meat was to be avoided (Clutton-Brock, 1999, pp. 178-182). During this time, domestic rabbits became larger than their wild ancestors, with a change in skull shape that included a larger face and smaller cranium. During the Middle Ages, rabbits were not bred for coat color. However, as they spread through the world, many different coat colors, patterns, and fur types were selected in the formation of different breeds. The wide distribution of domestic rabbits resulted in many regional varieties or breeds. There are Dutch, Polish, Flemish, and many other breeds associated with their lands of origin. Some of these breeds were selected and maintained for their meat and others for their fur. The New Zealand white rabbit is hugely popular as a very versatile breed. It is hardy, provides good quality meat and white fur that can be used as is or easily dyed, and has also been widely used as a subject in biomedical research. A variety of dwarf breeds have been developed for pet keeping. Rabbits were first brought to the New World in the 1700s as an easy and convenient animal to raise for food. The number of breeds present in the United States increased dramatically in the 1950s, when keeping rabbits became more popular as a hobby (Gendron, 2000).

Rabbits were originally classified as a type of rodent until 1912. They are now correctly placed into the order Lagomorpha with pikas and hares. Lagomorphs have a unique digestive system. They feed primarily on grasses and other fibers. They are unable to retain the full nutritive value of these foods after the first pass through their digestive system. Their feces after the first pass through the system are soft and moist. Bacteria found in their colons further break down the nutrients in the food. The rabbits then eat these first feces to digest and absorb additional nutrients, especially B vitamins. This behavior is known as coprophagy.

In the past 15-20 years the concepts of how to keep and care for companion rabbits has changed dramatically. Rabbits were generally kept in outdoor hutches, even when being kept as pets. They were rarely considered a house pet. The term house rabbit is now common and is indicative of this changing philosophy. Rabbits can be litter-trained and allowed supervised liberty in a home. Care must be taken with any houseplants that may be present and electrical wires (to which rabbits seem to have an unnatural attraction). They may be confined in a cage or hutch indoors when it is not possible to supervise their activities. However, this move indoors with freedom to move about is another step in the evolution of the rabbit (Figure 2-12). Rabbits have moved from the role of livestock as a source of food and fiber, to pets and objects of a hobby or fancy, to true companions for some people.

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Gerbils. Gerbils (Meriones unguiculatus) were first brought to the United States for research by Dr. Victor Schwerker in 1954. They became a popular alternative to hamsters once they entered the pet trade. Their long tails are furred, blunting some of the squeamishness that people have stemming from experiences with wild rats and mice. They are also more active in the daytime than hamsters and other rodents, providing more opportunity for interaction with people. Clark and Galef (1980) studied and compared domestic and wild gerbils. They found that the domesticated gerbils were less aggressive, had smaller adrenal glands, and grew more rapidly than the gerbils from wild population. These are all changes one would have expected as a result of domestication. In addition to these changes, breeders and fanciers have been able to identify and select for a variety of coat colors. Gerbils can be found in colors ranging from white albinos, to cream, fawn, and completely black (Figure 2-13).

Rats and Mice. Keeping rats and mice as pets challenges many cultural and personal aversions to rodents. Wild rats and mice have long been a scourge of humans. As soon as humans were capable of gathering or producing enough food to store some part of it for use at a later time, it is likely that rats and mice were finding ways into the storehouses to eat their fill and spoil additional quantities with their feces and urine. As noted earlier, cats initially found a welcome in the world of humans when they became allies in the battle against rodents. The ferret was another species that was enlisted to assist in this effort. Several of these highly adaptable rodents tagged along on ships and other forms of transportation to follow humans and quickly spread from their Asian origins to worldwide distribution. Three rodent species have established themselves as commensal species of humans (Clutton-Brock, 1999). The black rat (Rattus rattus) is a native of Asia Minor and probably spread through Southern Europe with the Romans. It is believed to be a significant carrier of many diseases. The brown rat (Rattus norvegicus) spread across Europe in the mid-sixteenth century. The brown rat is highly adaptive and prolific. It has a wide-ranging omnivorous diet and can withstand significant climate variation. The house mouse (Mus domesticus) probably spread through the world before the rat species and may be the second most populous mammalian species on earth after humans.

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Tamed mice were kept as pets for at least 300 years. In the mid-seventeenth century, white, grey, and dark varieties were noted. They became common subjects for biomedical research as the need grew for a small mammal that was easy to maintain and breed. Their short life cycle and prolific reproductive talents were well suited to the study of genetics. Gregor Mendel's research in genetics was rediscovered in the early 1900s, igniting a wave of research in the field that continues into this century. By 1909, Clarence Cook Little was producing inbred strains of mice, providing a significant advance and an important new tool for research. Genetically engineered strains of mice are now a significant resource for a wide range of research. The genetic diversity that made the mouse an ideal species for biomedical and genetics research also provided a rich substrate for fanciers interested in producing a variety of different types of mice. Pink-eyed, albino mice may be the form most familiar to most people. They are bred and sold in large numbers as pets, and as feeders for people who keep various reptiles as pets. Fancy mice that are bred and raised by hobbyists come in a wide range of colors, patterns, and several coat types (Figure 2-14). They are typically larger than the standard albino mouse. Coat colors may be creams, browns, blacks, in solids (or also called selfs), and various combinations in piebald and other patterns. Silky mice, with a fine, shiny coat, in gold or silver coat colors are particularly striking.

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Brown rats or Norwegian rats have also been domesticated. They have also been bred for use in the laboratory and as pets. The albino rat became a standard subject for a great many studies in both biomedical research and psychology. Their ubiquitous presence in the mid-twentieth century in American psychology laboratories led Frank Beach to publish a classic paper, with the wonderful title, "The Snark Was a Boojum," on the weakness of comparative research when the work was so highly dependent on a single species for study (Beach, 1950). Rats are still used in biomedical research, and many different genetic lines have developed for this purpose. Many of the domestic rats kept as pets are colored varieties. Similar to fancy mice, various blacks, browns, and combinations are popular color forms for rats kept as pets (Figure 2-15). Rats are generally neat and tidy. When socialized, they will be quite tame, and will recognize familiar people and willingly perch on their shoulders and accept treats.

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Guinea Pigs. The domestic guinea pig, or cavy (Cavia porcellus), is descended from the wild cavy of South America (Clutton-Brock, 1999). The guinea pig is one of the few New World species that was successfully domesticated. The Inca civilization in Peru raised guinea pigs for food. In villages there they may still be allowed to live within a home, and killed as needed for meals (Caras, 1996). Sailors returning from the New World would carry along a supply of cavies to provide fresh meat on the voyage home. Presumably, animals that made it back to the sailors' home ports alive might end up as intriguing pets for sweethearts and children. Their small size, chubby bodies, and habit of whistling would all seem to have predestined them to move from the pot to pet (Figure 2-16). The origin of their common name, guinea pig, is uncertain. One popular theory is that the ships returning to Europe frequently stopped in West African ports, including Guinea, and hence the name. Another theory is that when they first appeared in England they could be purchased for an English coin called a guinea. The "pig" part of the name is more than likely derived from their habit of squealing when aroused.

Hamsters. The origin of the common, or Syrian or golden hamster (Mesocricetus auratus), is well known (Clutton-Brock, 1999). In 1930, a single female hamster and her young were dug out of their burrow near Aleppo in Syria (Hemmer, 1990). Hamsters from this line were brought to England in 1931 and to America in 1938. They were initially used as laboratory subjects and later became popular as pets. A number of different varieties have been developed, including teddy bear with longer fur, and different coloring patterns (Figure 2-17). Several different, smaller, dwarf hamsters have entered the pet trade in the last fifteen years. The Russian dwarf (Phodopus campbelli), or Campbell's hamster, is among the most common of these. Others include the winter white (P. sungous), Chinese hamster (Cricetulus griseus), and Roborovski hamster (P. roborovskii), available less often in retail pet stores. It is striking that a nocturnal rodent with an occasionally pugnacious personality would become such a popular pet for children. However, the hamster is small and generally inexpensive. It has the striking habit of packing its cheeks with food and can be comical in its behavior. Just as important perhaps has been the ingenuity of the pet product industry in the development of a vast array of imaginative cages and environments for the hamsters. Plastic tubes connecting a variety of modules for sleeping, eating, and exercise have continued to stimulate interest in the animals. The monetary value (or lack!) of the actual hamster to the retail industry is reflected in sales where the purchase of a complete habitat may come with a free hamster to live in it.

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Livestock

A number of people raise and keep livestock as companions or as hobby. Many farm families hand-rear and raise orphans or runts, keeping them around as pets for the children. In the end, these pets may or may not end up going to market along with their brethren who had stayed with the flock or herd. The emotional dynamics of this process have stimulated quite a few children's books, including Charlotte's Web by E. B. White. The current growth in keeping livestock as a hobby is associated with an interest in unusual or rare breeds. Miniature horses are raised for show and will compete pulling small driving carriages and carts. They have even been bred and trained to fill the roles normally held by Seeing Eye or assistance dogs. Miniature cattle allow would-be ranchers to operate on small plots of land. They are also a curiosity that people enjoy seeing and petting (Figure 2-18).

The fanciers most serious about keeping livestock as a hobby may decide to concentrate on rare or endangered breeds. The development of modern production agricultural methods and a commodity-based economic structure has led to concentration on just a handful of breeds in each species. This has led to the loss of some older breeds that do not meet the needs of these high-volume production systems. The American Livestock Breeds Conservancy (ALBC) was formed in 1977 to conserve historic breeds and genetic diversity in livestock (see <http://albc-usa.org/>). These breeds include asses, cattle, goats, horses, sheep, pigs, rabbits, chickens, ducks, geese, and turkeys. They point out that these now-rare livestock breeds are an essential part of America's agricultural past and an important resource for the future. These breeds were developed to meet the needs of specific geographic regions and climates. Their genetic diversity may be a critical resource if agricultural changes require new adaptations in livestock.

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The ALBC maintains a conservation priority list of over 150 different breeds based on genetic and numerical parameters. These parameters include (poultry have separate requirements):

1. The breed is from one of the seven traditional U.S. livestock species mentioned above.

2. The breed census satisfies numerical guidelines such as

* Critical: Fewer than 200 annual registrations in the United States and estimated global population less than 2,000 (for rabbits less than 50 annual registrations and a global population under 500)

* Threatened: Fewer than 1,000 annual registrations in the United States and estimated global population less than 5,000 (for rabbits fewer than 100 annual registrations and a global population under 1,000)

* Watch: Fewer than 2,500 annual registrations in the United States and estimated global population less than 10,000 (for rabbits fewer than 200 annual registrations and a global population under 2,000); also included are breeds that present genetic or numerical concerns or have a limited geographic distribution

* Recovering: Breeds that were once listed in another category and have exceeded Watch category numbers but are still in need of monitoring

* Study: Breeds that are of genetic interest but either lack definition or lack genetic or historical documentation

3. The breed is a true genetic breed (when mated together, it produces the breed type).

4. The breed has an established and continuously breeding population in the United States since 1925. This requirement includes stipulations related to the number of breeding lines in the United States, the number of breeding females, and the presence of an association of breeders, among other specifics.

The Jacob sheep is one interesting and typical example of a breed listed as rare by the ALBC. They are a spotted sheep and descriptions date back to biblical accounts of Jacob, who bred spotted sheep. In the 1600s, spotted sheep were documented in England, and they became popular as an ornamental species. They are small and hardy, and have the interesting characteristic of being multihorned (Figure 2-19). The Jacob sheep were imported into the United States in the 1900s. They have grown in popularity as their fleece is favored by hand spinners and weavers practicing traditional crafts.

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In addition to hobby breeders, a number of historic villages and restorations have also begun to keep rare breeds of livestock on their grounds. The animals add to the pastoral setting, providing a glimpse of the intimate relationship that colonists had with the animals they kept. Maintaining the rare breeds is also consistent with their efforts to preserve and exhibit the culture of the era they are trying to represent.

DISCUSSION QUESTIONS

1. What is the relationship between domestication and tameness?

2. How does the reproductive potential of a domesticated species typically compare with that of its wild counterparts? Why does this occur? What are specific changes that occur?

3. What is neoteny? Besides those described in the text, give an example of physical neoteny and behavioral neoteny.

4. What are the limitations of using archaeological evidence to study the relation of dogs to wolves?

5. Compare and contrast the traditional view of canine domestication (i.e., wolf taming) with the alternative process involving the food source provided by human civilizations.

6. What morphological changes accompanied the increase in tameness of silver foxes observed in the classic study by Dmitry Belyaev and colleagues?

7. Describe the timing of and motivation behind the development of early dog breeds. What were the characteristics and purposes of the first breed types?

8. From the Middle Ages to the present, describe the spectrum of care received by dogs according to the socioeconomic status of their people.

9. Describe the development of kennel clubs and the dog fancy. What role has this process played in the establishment of today's puppy mills?

10. Describe the incorporation over time of cats into human civilizations. How did the cat's domestication differ from that of the dog?

11. How did the treatment and perception of cats during the Middle Ages differ from previous times? What factors were involved in this change?

12. Compare and contrast the development of the cat fancy and the dog fancy. Did cat breeds develop for the same reasons as dog breeds?

13. How has the role of the horse changed throughout history? What roles do horses play in modern American society?

14. Many of the small animals currently kept as pets were originally used for functions other than companionship. Select three of the small animals discussed in this chapter and describe their original functions. Are the animals still used for these purposes today?

15. How has domestication affected the coat color of small animals? Are similar effects observed in the cat and dog?

16. What is the goal of the American Livestock Breeds Conservancy (ALBC)? What led to its development? What types of animals are involved?

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Author:Zawistowski, Stephen
Publication:Companion Animals in Society
Geographic Code:1USA
Date:Jan 1, 2008
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