Dogs: A Startling New Understanding of Canine Origin, Behavior & Evolution

Dogs: A Startling New Understanding of Canine Origin, Behavior & Evolution

by Raymond Coppinger, Lorna Coppinger




Marking the first time that dogs have been explained in such detail by eminent researchers, Dogs is a work of wide appeal, as absorbing as it is enlightening.

Drawing on insight gleaned from forty-five years of raising, training, and studying the behaviors of dogs worldwide, Lorna and Raymond Coppinger explore the fascinating processes by which dog breeds have evolved into their unique shapes and behaviors. Concentrating on five types of dogs—modern household dogs, village dogs, livestock-guarding dogs, sled dogs, and herding dogs—the Coppingers, internationally recognized canine ethologists and consummate dog lovers, examine our canine companions from a unique biological viewpoint. Dogs clearly points the way for dog lovers, dog therapists, veterinarians, and all others who deal with dogs to understand their animals from a fresh perspective.

How did the domestic dog become a distinct species from the wolf? Why do different breeds behave differently? Most important, how can we improve the relationship between humans and dogs?

The authors show how dogs' different abilities depend upon the confluence of their nature and nurture—that both genetics and the environment play equally key roles. They also reveal that many people inadvertently harm their canine companions because they fail to understand dogs' biological needs and dispositions.

Dogs is a highly readable biological approach by noted researchers that provides a wealth of new information about the interaction of nature and nurture, and demonstrates how unique dog behavior is in the animal world.

Product Details

ISBN-13: 9780684855301
Publisher: Scribner
Publication date: 05/28/2001
Series: Lisa Drew Books
Pages: 352
Product dimensions: 6.12(w) x 9.25(h) x 1.20(d)

About the Author

Raymond Coppinger is professor of biology at Hampshire College and the author of Fishing Dogs. A former sled dog racing champion, he now lectures widely about dogs.

Read an Excerpt

Chapter 7: Household Dogs

Working dogs may enjoy a mutualistic relationship with people. But once evolved into a working shape and behavior, these dogs also get adopted as pets. And here lies a dark side to the association of dogs with people. Just what is the ecological status of people and household dogs? Household dogs — also known as pets or companion animals — are usually considered to be beneficial for people, providing unquestioned love, constant loyalty, eager companionship, and a variety of what people perceive to be positive additions to their lives. But in the scheme of biological survival and perpetuation of the species — any species — what counts is not what we perceive to be beneficial, but what really is beneficial, biologically. For living things, survival depends on a trio of absolutely fundamental needs: food, safety, and reproduction. Unless dogs provide measurable quantities of these essentials, they are not strictly beneficial to the survival of humans.

At the same time, people are usually considered beneficial for dogs, furnishing their food, safety, health care, jobs, and, often, well-arranged opportunities to reproduce themselves. However, as a biologist looking closely at dogs, I become more and more uneasy. At the very least, I think the symbiotic relationship of pet dogs with their owners is seriously unclear. Perhaps that is because there is more than one association between them. Part of the pet-owner relationship is mutual, in the direction of the working dog and its owner. But sometimes I see the population of pet dogs acting like parasites on people. And at other times I see humans treating dogs really badly.

Maybe people don't knowingly treat dogs badly. It could easily be that pet owners are just not biologically good for their pets, sustaining a relationship that ecologists call amensalism.

Amensalism describes the biological relationship in which one species is not affected by the association, but the other species, by accident, is hurt by it. An ecology book describes the relationship between bison and prairie chickens as amensalism. Prairie chickens live in bison country, not really affecting the bison, but as the bison search for food they step on prairie chicken eggs. They don't mean to, but nevertheless they are bad for the birds. Amensalism is usually contrasted with parasitism, in which the parasite purposely lives off the host and saps the host's strength.

Can I cite an example of amensalism? Easily. Take just one case — that of the bulldog. From active and noteworthy employment as the butcher's working dog, and then as a sporting dog, the breed has been adopted as a pet and show dog. In order to enhance its robust, highly unusual appearance, breeders have selected for those traits that emphasize the essence of bulldog — the thick, massive head and short, pug nose. What they have achieved, probably accidentally, are dogs that often can barely breathe, can barely chew, whose puppies are hard to deliver, and females that have to be artificially inseminated. Such animals cannot be living a comfortable life. Their "enhanced" abnormal shape traps them in a genetic dead end. Being caught and bred as household dogs is detrimental to their long-term reproductive survival.

Likewise, I can easily cite an example of the dog being a parasite on humans. Parasites do not usually kill their host, but they do decrease the fitness of the host, and reduce the energy available to the host for its own survival. The two species live together, and one gets its food at the expense of the other. In its simplest form, I go to work, earn a salary, and part of my salary is spent on dog food to feed my pet dog. At its worst, my dog has a disease or an injury that costs me hundreds of dollars to try to cure. Or, it bites someone and I get sued. Or, the dog has behavioral problems and ruins furniture. The dog is a drain on my resources and my energy. The dog takes time and money that I should be investing in my children. It makes me less fit for survival.

The fifty-two million dogs living in households in the United States were probably added to those households as the result of a conscious decision by the householder. People ask me, "What kind of dog should we get for a pet?" What they are really saying is "What kind of dog do you think would benefit us the most?" Often the questioners have been thinking purebred dog. It could be the first time they have purchased a dog, and many people consult friends, relatives, books, and the occasional dog expert for advice.

I often start my answer with "Whatever you decide, get a pup before it is eight weeks old, and spend a lot of time with it during the next eight weeks." I guess my assumption is that if they ask the "what kind of dog" question, they think a breed is a package of behaviors that comes prearranged. They are usually unaware that any and all puppies need an informed and thoughtful owner to shape the pup's course toward well-behaved adulthood. Their assumption is that the right breed is all that is needed to effect the perfect dog-human bond.

They hardly ever ask me, "If you were getting a dog, what would you get?" Sometimes when I suggest breed x, they immediately say, "Oh! But we wanted a big dog." And I ask, "Why would a big dog be more of a benefit?" Are they looking for protection? The conversation suggests that big dogs have more of an aesthetic impact. The big dog enhances their image of themselves. When they say big dog, I just say, "I don't know the answer." I can't fathom a big dog as a companion. I want something that could go with me everywhere in the car. I'd prefer a small dog with a smooth, dry coat, and no long tail to get slammed in the door by accident.

On an estate in central England, I once interviewed Lady Richards, whose dogs enjoy expanses of greensward, intriguing woodland copses, and a peaceful pond. She had a lithe and leggy lurcher (hers was a cross between a greyhound and a border collie) that she was very fond of. She said, "We are fortunate in that we can afford to entertain such a dog." She had the right idea: What is it that I as a pet owner possess that will enhance the dog's life?

The relationship should not be a one-way street, where I'll get a dog that pleases me, and if it continues to please me, I won't turn it in to the local shelter.

My dilemma about which symbiotic relationship operates between pets and people is slowly resolving. Especially when I imagine a person purchasing a ten-week-old puppy, locking it in his apartment while he goes to work, coming home to find the apartment trashed, consulting an expensive dog psychologist who after many months says it is hopeless, and turning the dog in to the local shelter, which euthanizes it. Bad situation for the person, disaster for the dog.

Household dogs are what I think of as family pets. I understand there are many variations on the theme. There is even discussion about whether household dogs should be called companions rather than pets. Pets, it is argued, are animals like goldfish or caged birds or snakes. Pets can be exotic or not: the Pet Rock was fashionable a few years ago. Joshua Slocum, on his single-handed sail around the world, had a pet goat for company. But "for company" doesn't necessarily mean companion. I think of a companion as different from a pet. A companion dog accompanies me during some of my daily activities, or shares with me some task, like pretending to be a coyote so I can evaluate the response of a guardian dog.

My border collie Jane was a companion, not a pet. I bought her and her brother as pups from Will Wilson in Scotland in 1977. Jane participated in all our herding-dog experiments, lived in the dorms with my students, and traveled with me everywhere. She was about coyote-sized, and had pricked ears and a sharp muzzle like a coyote. I would use her as a mock predator. I'd send her into a flock of sheep to test the effectiveness of the livestock-guarding dogs. More than once she had to break her outrun and turn tail for me, outracing the guardian dog and at the last minute leaping into my arms, to be cradled and praised. She was also great fun at lectures and demonstrations. It was easy to get her to show the predatory motor patterns. She could not take her eyes off the ball waving around in my hand and the audience would be in stitches. If I put the ball on the floor Jane would eye-stalk it, and if it didn't move she would do a perfect mouse jump on it to get it to move. She joined me on numerous television programs to herd sheep and show the differences in motor patterns between herding and guarding dogs. I used her once in class to demonstrate cognitive differences between children and dogs. From a very young age children understand the pointing finger in ways dogs — even Jane — cannot.

Jane lived in my van, and therefore I spent some part of each day with her. When I was on the road during the years of the livestock-guarding-dog project, we spent twenty-four hours a day together, even sleeping in the van in remote pastures. Although we were constant companions, I didn't really like her. She was an annoying, high-strung dog; she was driven, absolutely driven to display those internally motivated herding-dog motor patterns. She'd stand at the long side window of the van wiping her nose against it as she eye-stalked passing cars. I'd find myself yelling at her to "Git down! Git off my bed!" and milliseconds later she'd be right back. I built her a padded box but she'd start wailing to get out. She never stopped moving. She was always in my face or my ears. It was awful. Would I get a border collie as a pet? Goodness no! It was bad enough having one as a companion.

If a novice dog owner got Jane as a pet, he'd have taken her to Nick Dodman's (The Dog Who Loved Too Much) office at the School of Veterinary Medicine at Tufts University for a big dose of tranquilizers. I would have staked my reputation that any of the operant-conditioning specialists would have wound up seeking counseling themselves after a week with Jane. At obedience school, a professional trainer ended up holding Jane at arm's length (while she chewed on it), yelling, "You leave this dog with me, and I'll have her trained by Monday!" I was tempted, but I liked the guy and wouldn't have let him destroy himself attempting to modify Jane into a submissive dog.

Jane illustrates the difference between companion and pet. But I'll call them all household dogs, which is more of an ecological definition. The role of pets, I believe, is the muddiest of relationships between human and dog. I think it is also the least understood in terms of any benefits of the relationship. And, most serious of all, I think the greatest abuses to dogs occur in this home-dwelling population.

Ecologically speaking, as I noted in the Introduction, the domestic dog is an incredibly successful species. Populations of animals grow from small beginnings, colonize the available niche, and at some point reach equilibrium with their environment. As long as the habitat sustains their ability to find food, avoid hazards, and reproduce, dogs do very well.

Right now the population of household dogs in the United States seems to be fairly level. The pet figure may have stabilized because the human population has stabilized, or it may be that people are getting wise and beginning to find out that household dogs are not worth the expense.

Here at the beginning of the twenty-first century, over fifty million household dogs live in the United States. Europe houses an estimated thirty-five million. Dogs in Europe are allowed and routinely appear in restaurants and public buildings, on trains and buses. My experience with friends in Europe may not be typical, because I tend to spend time with people who have dogs. All I know is I have trouble fitting my legs under the restaurant table because of the dogs. As a result, I tend to think the thirty-five million figure is low. If I add Canadian dogs to these populations, I get one hundred million household dogs in the industrial West.

In the United States each year, households produce 3,700,000 puppies. Hobby breeders produce another two million, and half a million are produced by commercial breeders for department store and other retail sales. That is a turnover of 6,200,000 dogs a year. If the population is not going up or down, then 6,200,000 dogs die every year. That is a 12 percent annual mortality rate, which for a species with a life span of a little over ten years is a low mortality rate in the wild.

In the United States, four million of these dogs spend part of a year in animal shelters. For 2,400,000 of them it is the last stop. Almost 5 percent of our companion animals are dogs nobody wants, and they get "put to sleep." Culled. Again, disaster for the individual dog. Some of this culling may be related to competition between people and dogs for food resources. People soon decide they can't afford the dog, and turn them over to humane societies.

What is the biological relationship that fosters the symbiosis between household dogs and people? I'd like to look first at the case for parasitism. If dogs are parasites, and they exist in the United States at the ratio of 1.5 dogs per dog-owning household, then as a human being I am more than a little concerned about this relationship. Among the costs dogs impose on humans are many direct expenses: food, veterinary care, and management (collars, licenses, fences). Dogs also have an impact on society's budgets in the fields of health care, sanitation and medicine, insurance, legislation, and law and order. They act as vectors for disease and as serious nuisances due to incessant barking or destruction of property.

Behavioral ecologists almost always start a research investigation with information on a species' food supply. Dog food is manufactured by companies that purchase the ingredients from the wholesalers of grains and meat products which are also used in human food. They buy on the same auction blocks as do the processors of human foods. Household-dog food ingredients are not leftovers or waste products. The companies will aver that the dog food is not the same quality as human food, that the grains are not good enough to be milled for human food. Some will claim that the animal products are offal and other by-products not edible by people. Sometimes they just say it is surplus human food.

Dog food companies have one big, limiting problem: they cannot frequently or substantially change the formulation of the food. Even tiny changes in ingredients or processing give dogs digestive upsets. A dog eating the same food day after day gets accustomed to that formula. All the little microorganisms living in their digestive tracts are in ecological balance with the incoming ration. Change any of the ratios, and the microbes begin warring with each other, which shows up as diarrhea. This is the reason why changing brands must be done slowly and gradually — so the dog can adapt to the new ration.

Because of this, many people will change brands if their dog's digestion is disordered. Dog food companies are very sensitive to that, not wanting to alienate a loyal customer from his favorite brand. Thus, every day of every week of every month of every year, they have to be buying not only the same ingredients, but the same quality of ingredients. They are not searching around for the lowest-quality waste products. They are locked into obtaining a consistent quality and constant supply. The added cost of buying good raw materials is a minor part of the overall cost of finding the tons of edible materials, and then the manufacturing and distribution of them. Probably the advertising costs more than the ingredients.

Dog food is edible and nutritious for humans, and some people count on it being perfectly safe to eat. (I am not recommending this.) There are reports that in some city districts, more dog food is sold than could possibly be eaten by the estimated dog population. The assumption is that poor people are feeding it to their families. Questions about putting antibiotics or antihelminthics and other medicine in dog foods have to be judged on the probability of its being harmful to humans, because sooner or later some of the dog food ends up in human mouths.

I understand that farmers don't grow food for people, they grow food for money. They grow wheat or beef because it is saleable, not because it is human food. Farming is the process of people turning sunlight into food on farmland. But what is important here is that we in Western cultures have achieved an advanced form of Neolithism, that is, farming with technically advanced tools. All but a tiny fraction of our food is commercially produced by farmers. And our food base is the same as the pet dog's food base. Dog food is not waste products. Therefore its production does have ecological consequence.

Consider this. A dog's normal body temperature is 101.5 degrees Fahrenheit; a human's is 98.6. That three extra degrees means it takes more calories to maintain each pound of dog. In fact, given certain conditions of size and activity, and ambient temperature, dog cells can require as much as twice as many calories as human cells — all other things being equal, which of course they never are. Rising body temperature is not a linear progression. It takes many more calories to raise body temperature from 101 to 102 degrees than it takes to raise it from 96 to 97 degrees. Reports from the Iditarod indicate that a fifty-pound racing dog ingests up to ten thousand calories a day. Several Iditarod champions feed their dogs many times a day during the race in order to avoid the physical loading and digesting problems of feeding ten thousand calories all at once.

Comparing a population of dogs to people would illustrate how expensive dogs are. Assume the average household dog weighs twenty-five pounds, about the size of a small beagle or the Pemba hound (the generic dog). Since many of our most popular breeds are German shepherds and Labrador and golden retrievers, which weigh about seventy pounds, twenty-five pounds seems a low estimate for my illustration. But over six million of these dogs are growing pups less than a year old. So the twenty-five-pound figure might actually be a little high.

I'll assume that the average American weighs one hundred pounds. If dogs need twice as much food per pound as humans, then fifty pounds of dog eats as much as one hundred pounds of human, or two dogs eat as much as one American. Thus, the fifty-two million pet dogs eat as much as twenty-six million people. American dogs eat as much farmer-grown food each day as all the people in New York, Chicago, and Los Angeles combined.

Everybody I tell this to — my friends, veterinarians, biologist colleagues — reacts to that figure as a huge exaggeration. "It can't be!" they say. But I don't think it is much of an exaggeration. It is a tough figure to calculate. The calories needed per pound of dog decrease as the weight of the dog increases. Great, big, reasonably inactive dogs don't require many more calories per pound than a hundred-pound inactive person. But twenty-five-pound inactive dogs require 1.5 times as many. And activity requires more calories. When I add the requirements of growing puppies or children, the calorie needs jump to two to three times the resting figures. Keep the dog outside in the winter and the figures jump again. Even if you wanted to ignore the extra three degrees of a dog's temperature, and consider dogs energetically equivalent to humans, they would still require as much food per day as all the people in New York, Boston, and San Francisco. This for me is still an astronomical figure.

Imagine being responsible for having enough land and growing enough food to feed all the people in three major American cities every day. No wonder the dog food business is such a great idea. Using the back of the bag of my favorite dog food to figure the costs involved, I find that it recommends I feed my twenty-five-pound dog 225 pounds of food a year. At 40 cents a pound, the cost for all the dogs in the United States is in the billions of dollars.

There are other ways of calculating the costs of feeding fifty-two million dogs. I once visited a communal farm in China where the claim was that four hundred people were getting all the food they needed from 150 acres of land. (They were later severely criticized for exaggerating.) That is more than a third of an acre per person, to meet all their nutritional requirements. (In the United States we farm 3.5 acres for every person, but a huge amount gets sold out of the country and we stockpile surpluses, so I don't know how many acres it actually takes to feed us.) The Chinese brigade was growing enough food to feed eight hundred American dogs, or about five dogs per acre. Translate that statistic to feeding American dogs, and it takes fifteen thousand square miles of farmland to feed our fifty-two million pet dogs. (That, by the way, is "only" one percent of our farmland). That is an area as big as Massachusetts, Connecticut, and Rhode Island combined. An area approximately the size of Denmark or Switzerland or Taiwan.

That is also an area four times the size of Yellowstone National Park. Of course, Yellowstone is not prime agricultural land. If we took the fifteen thousand square miles of prime farmland and turned it into national park, we could have the greatest wildlife sanctuary in the world. Imagine the number of wild things and wild habitat that are displaced by the necessity of growing dog food.

So far all we've done in this discussion is feed the household dogs. We haven't considered the pollution from their feces, which has to be equal to or worse than twenty-six million people eliminating al fresco. Neither have I added the medical cost of dog bites, which some experts think are in epidemic proportions. My home state of Massachusetts averages twenty-one thousand dog bites per year that need medical attention. The post office runs a course for employees on how not to get bitten. Society often demands that dangerous dogs be culled. England has dangerous-dog laws, which appear as an almost desperate attempt to protect people from dog bites. And, of course, some few people do get killed by a dog that purposely sets out to kill them. I talked to a man who hated dogs because one pulled his daughter off a bicycle, and when she hit her head she lost an eye.

Some costs are incalculable. How do you compute the cost of lost sleep from the dog barking next door, the cost of running animal shelters, the cost to police forces of finding and returning lost dogs? Most towns have a dog officer who spends all or part of his time on dog problems. Humane societies sponsor conferences for experts who try to establish policy and write intelligent laws about how to protect ourselves from dogs.

Where does all this fit on our symbiotic scale? Is this an example of commensalism or mutualism? The commensal village dogs scrounge the wastes of human foraging. Like rats, they eat what the humans cannot. But they also carry diseases and bark at night. It doesn't matter much to people that they are there, because they exert neither major benefit nor major cost. The mutual working dog performs a service for its food. But household dogs are not only eating at the same table as people, but from the same plate.

Household dogs are competitive for our food and other resources. Their numbers may not have reached the point where this competition seriously collides with our ability to feed ourselves, but that is no reason to ignore the facts. In addition to being well fed, dogs get treated like family members in many other respects. At the same time, they can be in direct competition with human children for the time and resources of the parents. They can inflict substantial harm. If I were a brave ecologist I'd say the household dog sounds like a parasite.

The only argument that would convince an ecologist that household dogs are not parasites would be to show benefits to people for investing the level of resources we do into that population. The benefits would have to outweigh the costs. And I doubt that anybody could do that.


The purported benefits of dogs usually fall into two categories: 1) they work for people, providing a tangible service, and 2) they make people feel better in some meaningful way. The category here is household dogs. The subject is pets and companions, not working dogs. The fact that generations ago individuals of a breed were sheepdogs, hunting dogs, or police dogs does not mean that this particular household dog could, even if trained, perform those tasks. Neither does it mean any of those tasks would be useful around the house. In fact, many of the phone calls I get about dogs are from people complaining about an overzealous working dog. And this is absolutely true: while writing this very paragraph, someone called offering me a well-bred Queensland blue heeler — overactive in its breed-typical motor pattern displays, and unbearably obnoxious with its heel-nipping. It was driving its owner nuts.

Among the tangible benefits commonly claimed for household dogs is protectiveness — they are watchdogs. Such dogs supposedly react appropriately to hazards, thereby increasing the hazard-avoidance abilities of the owners. As far as I know there are no data that people with dogs survive more fires, have fewer burglaries, get mugged less, or are more often alerted to other household catastrophes than people without dogs. But the anecdotal evidence always favors the dog. The stories of heroic feats by dogs attract human interest. Cynics call them "Lassie stories." They are stories of dogs that find their way home over hundreds of miles, or perform some insightful act requiring cognitive skills on the order of human intelligence. It is always unpopular to question such miracles. We all want to amplify our relationship with dogs.

But in my experience the average dog can barely find its way home from next door. My cousin Barry's retriever, Rosie, got lost daily chasing cars. She'd start out in front of the house, chase the car to the corner, and become disoriented. I've searched for so many lost dogs I vow each time I do it that I will never do it again. Police blotters have columns of lost dogs and columns of found dogs, and there is seldom a match. Our local radio station has daily lost-dog reports, which they run as a community service.

Dogs don't run into burning buildings, nor are they capable of pulling unconscious people out of one. More likely, the dog knocks a lamp over and starts the fire. The one firsthand experience I ever had with a "heroic" dog was a few months ago when my fishing buddy and I rescued a fellow whose boat had capsized in a fast-running tidal river. He would have been fine clinging to his boat, except his dog panicked and, in trying to scramble up onto the overturned boat, she kept scrambling up her master's back, pushing him underneath the water repeatedly. By the time we got there he was so exhausted he was going down, on the way to being drowned by his own dog. The "heroic" part of the story is that his calls for help were so weak, I'd never have seen him in the dim light except I kept seeing his dog's head come high out of the water, and went to investigate the phenomenon. The dog turned out to be of a breed uncommon in New England, a Catahoula leopard cow-hog dog, which we also rescued.

For me to be cynical about dogs would be incongruous. Of course there are true, unaugmented stories of lifesaving by household dogs. For anyone who has experienced a dog's ability to avert disaster, there is no question whatsoever about the value of the dog in society. And naturally I am well aware of the accomplishments of guide dogs for the blind, of dogs that assist physically challenged people in their daily lives, of dogs used in search-and-rescue and police work. The sight of a well-trained dog performing a task that would be difficult if not impossible for a human is special in the animal world.

In the last two decades of the twentieth century, the status of dogs in relation to people attracted scholarly study and led to the documentation of certain benefits of dogs to the psychological and medical well-being of people. There is much more to this category than buying a dog for a child in order to teach empathy and to care for other living things. I must admit, however, that my motive in acquiring a Chesapeake Bay retriever for our eleven-year-old son was to develop a career opportunity for him. Tim wanted to be a baseball player when he grew up, but since he lacked baseball-playing siblings or friends who would play out of season, he needed dogs to play the outfield. Scoter and my companion dog, Jane, used to retrieve fly balls for him all afternoon. The benefit was not Tim's, perhaps, but mine, since without the dogs I would have had to perform that function. Another benefit for me was that Jane would be so tired at the end that her obnoxious behaviors would subside for a short while.

The psychological benefits dogs can impart to people are measured in both precise and imprecise terms. Psychologists, biologists, and doctors have been measuring vital signs such as blood pressure, documenting longevity, and recording personal perceptions of people in critical circumstances — both with and without a dog. The results in favor of dogs are clear in many cases. Blood pressures in a wide array of social situations, among children as well as adults, are lower. Children are less apprehensive of doctors if there is a dog in the office. People who have a dog to return to after their first heart attack have a better life expectancy than the dogless controls. People with dogs have more access to certain social encounters. It is easier to talk to a stranger who has a dog. Strangers are more likely to talk to you if you have a dog. Walking the dog is a good way to meet other people who are also walking their dog. People confined to wheelchairs find the presence of a dog gains them positive attention from non-wheelchair-bound passersby, which has a positive social effect.

There are good data that say people in wheelchairs with a trained dog feel more like traveling in public. The dog enables them to engage in activities they otherwise would avoid. Blind people often specifically request German shepherd dogs as guides because they feel the dog would confer protection as well as guidance. Even if there isn't a ghost of a chance the dog could cope with a wrongdoer, the mere presence of the "police" dog enables these people to comfortably sally forth. Many people get a specific breed of dog for the home, or to jog with, that they feel would protect them. And if they sleep better because they think the dog is on duty, the dog must be construed as a benefit.

For the behavioral ecologist, however, the problem is to balance the equation. How do you measure the value of these benefits in terms of increasing the fitness of the human population? Old people living longer in a nursing home may perhaps be competing for their grandchildren's funds and their children's time. Spending time taking care of an aging parent past reproductive age at the expense of children at the beginning of theirs may not be to the species' biological benefit.

Again, however the benefit to an individual is construed, it has to be balanced against the costs to the society. The fact that less than one percent of blind people have a guide dog does little to offset the thousands of people each year who get bitten by household dogs.

From the dog's point of view, is this a good relationship to be in with humans?

My own belief is that the dog is not an ecological parasite in the true sense, even if the equation doesn't balance. A parasite evolves to the niche. A parasite benefits from its relationship with its host. Just as a cow is adapted to eating grass, a parasite is adapted to eating its host. Even though the dog may cost the human population, it isn't gaining much biological benefit from its household association with people. In fact, the system people use to propagate dogs may doom those dogs trapped in that system.

Therefore, I think the relationship that fosters the symbiosis between household dogs and people is amensal. Like the prairie chickens whose eggs are stepped on inadvertently by the bison, dogs are stepped on and hurt by people. Although people may believe they are caring for and "improving" their dogs, I'd like to approach their situation from the dog's point of view.

Dogs may appear childlike to some people, releasing caregiving behaviors. It has been postulated that one of the reasons dogs' heads appear more rounded and shorter-nosed than those of wolves is because people selected dogs to look puppylike, which is innately cute to people. Infant qualities also reduce aggressive interactions. But if short, rounded, puppylike dog faces benefit humans by giving them a release for child-rearing motivations, it is our preference that is being imposed on dogs. It is people who breed the household dog with the short face for their own benefit, without regard for the effect on the dog. Just because the human gets immense pleasure from dispensing care and affection on a dog does not mean that the permanently juvenile dog gets any pleasure or benefit from the short muzzle.

In the literature I read on human-animal relationships, the analysis of benefit is almost exclusively in human terms. Dogs make people happy, lower people's heart rates, help people survive heart attacks, increase the quality of life in nursing homes, help disabled and interned people have a better quality of life. Because dogs benefit people does not mean that people benefit dogs, however.

Just because dogs make "ideal" pets does not mean we should make them pets. Just because they have lovely, adaptable social natures does not mean we can exploit them with impunity. And just because they have infinitely malleable shapes does not give us the right to select for any deformation of the basic shape just because a different shape pleases us.

I think dog ownership has reached the point where humans have imposed on the good nature of dogs a little too much. I think that although those interested in dog welfare have done well to focus on the plight of dogs used in biomedical research — the so-called laboratory dogs — and to some extent the working dogs, they are ignoring a far greater and more insidious danger. I think far worse cruelty takes place with purebred dogs, adopted into the pet-class world, and in far greater numbers. I think the household-dog industry has taken too little responsibility for the consequences of present breeding policies and dog ownership.

The production and distribution of household dogs follows fairly standard methods. First of all, household pets are captured animals. "What kind of dog should I get?" says it all. These are not animals that adopt us, as the Pemban dogs do. Household fish in an aquarium are collected from the wild. The parent stock continues to breed in the wild. If the system works well, collectors take only a few "surplus" animals for the pet trade and basically the wild stock continues to reproduce on its own. The beautiful colors and the interesting shapes that we enjoy in our fish tanks are products of natural selection and subject to all the Darwinian rules of survival of the fittest. The stock remaining in the wild stays genetically fit.

It must be remembered, however, that the individual fish removed to aquariums are deprived of their reproductive rights and their genetic potential by our desire to enjoy viewing them. They are effectively reproductively removed from ever leaving offspring — they are genetically dead.

Household dogs are similarly captured. But they come from three different sources. Originally, over the millennia, people captured pups for pets from the (wild) village dogs. Like the aquarium fish, the parental scavenging stock continued to survive and reproduce in its original form. The farmyard dog and my dog Smoky's mother still have reproductive access to a large population of dogs, and are, like the reef fish, giving up a few surplus animals for human benefit. Similarly, capturing a livestock-guarding dog from the "wild" population on an annual transhumance also leaves the wild population intact and with access to a large population of dogs. This situation continuously maintains the genetic health of the individuals. In each of these cases the dogs are still responsible for reproduction, and their normal breeding habits and competition for limited resources will assure healthy offspring.

The second source of household dogs is retired working dogs. Jane was a working dog for me. She got old, and we thought she would be cold in the van at night and brought her in. She became a household dog in her old age. One could imagine hundreds of different scenarios of how a truly good working dog might end up in a home. In fact, the home might just be the most convenient place to keep the dog while it is not working. Many a hunter keeps his bird dog in the house, and on occasion I would bring a sled dog in. There might be advantages to having dogs inside. There's always the possibility of warmth: I've read that Eskimos and Australian Aborigines measure how cold it is by how many dogs it takes to keep them warm overnight.

But the breeding of these dogs is by artificial selection, and perhaps the problem starts here. The modern working-dog breeder breeds dogs that are good performers. I bred Jane just once. She was an awful mother who couldn't tell the difference between her pups and rats. She would eye-stalk them and forefoot-stab them to death if they moved. Here, selection for the eye-stalk behavior had created a biological monster. And I'm afraid that is the risk we run every time we start practicing canine eugenics.

The third source of household dogs is where the problems begin to mount. This is where the householder captures a working dog (breed) for a pet. And then the absolutely worst-case scenario, the breeding of working dogs for the household-pet market. Working dogs should not be pets. Working dogs should not be sold or purchased as pets. And working dogs should not be bred for the pet class of dogs.

If a dog is bred for exaggerated behavioral conformation and is expected to display it in a working environment, it is hard to imagine that the household environment is going to provide the proper stimulation for such displays. Often the person raising the dog has no idea of the critical period of social development or the specific requirements necessary to evoke the proper behavior. This results in dogs that have motor pattern displays not only inappropriate in the household environment, but that also can turn into compulsive disorders. A highly bred working dog raised in a nonworking household environment will still show the working behaviors it has been selected to display, but it will display them abnormally. Worse, it will display those behaviors in bizarre and obnoxious ways. My Queensland blue heeler would nip the heels of joggers. My best lead dog was a border collie that chased cars all day on a suburban street — which was why I got him.

Most working dogs are corrupted, if not ruined, for their job if kept as pets. How can a livestock-guarding dog protect livestock if it is in the house? Many working dogs transfer their socially developed protective behaviors in ways that make them dangerous to people. I avoided passing my guardian dogs on to people who didn't have livestock and wanted them for pets. I succumbed a few times, and I always thought it turned out badly, especially for the dog.

What is the difference between adopting a village dog or a working dog as a pet? The village dog is selected to have a low profile around humans. The village dog tries to maintain contact with the human, which is the source of food. The village dog is selected to solicit care from humans and not to threaten them. Thus, the difference between a village dog that grows up naturally as a scavenger of human resources, and one that is captured as a puppy and raised as a household pet is not all that great. In some cases, as in Pemba, I couldn't tell if a dog was a pet or not, and was totally surprised to find that the vast majority of them were not pets. "Is that your dog?" I asked the girl on Pemba, and it was difficult to discern from the answer what kind of relationship she had with this yard dog. Was it a pet, or was it just a cute scavenger that was protecting her yard from other scavengers? In these societies the difference between casting aside the waste and feeding the dog is a state of mind, an intention, rather than different behaviors.

Adopting an occasional village dog as a pet probably doesn't change the evolutionary dynamics of the wild population very much. The individual dog, of course, loses its place in the wild population and therefore may lose some reproductive opportunities. But it's more complicated for the working dog that is adopted away from its working environment.

The village dog is preadapted behaviorally to be a good pet. The purebred working dog is selected to show a variety of motor patterns that are often obnoxious in a household. Jane wiping her nose on my van window as she eye-stalked passing cars is a perfect example. Her internally motivated patterns (compulsive behavior) are also internally rewarded, and thus practically impossible to extinguish.

The question might be posed as to why one would want a working dog as a pet. Dogs like Jane are limited in their ability to adapt as perfect pets because of their innate behaviors, and thus perhaps they are sought because of the way they look. Something about the way the dog looks benefits the person. Perhaps a sled dog, even though it never pulls a sled, suggests that the owner is associated with the historical legacy or reputation of sled dogs. Perhaps the image evoked by the working dog enhances the owner's status or enables an association with other people who identify with the dog's heritage. Owning a hunting breed suggests that the owner might be a hunter, or might know about hunting and be a self-reliant outdoors person. I often wonder if I had lived in the suburbs whether I would have acquired racing sled dogs. It's hard to imagine they would have been welcome for very long, what with their daily howling and overwhelming cacophony whenever I arrived with the harnesses to hitch up for a training run.

Here is the big shift for dogs: no longer are they chosen for the way they behave, but for the way they look. Selection in the Darwinian sense is for their appearance. The benefit to the human is not in the innate behavioral abilities, but in the coat color, ear carriage, and size. But these are superficial traits, related to survival only cosmetically. It is ironic that the village dog, well suited to survive, is rejected as "just a mutt" for those same traits.

"What kind of dog should I get?" (to enhance my status...round out my image...amuse me...) does not consider the needs of the dog. It is based on the faulty assumption that any breed of dog is adaptable to the household environment.

During the past one hundred years, hobby breeders have taken the working-sporting breeds and bred them specifically for the household market. I understand that throughout history breeders have bred miniature and gargantuan forms of dogs simply for display: the bonsai-garden type of breeding. But few of our modern household breeds are much older than a hundred years. The "perfection" of breeds is coincidental with the interest in expositions in which owners or trainers submit their dogs to judges who decide which dogs are superior in looks. Over the past hundred years, the hobby breeding program has succeeded quite well in isolating subpopulations of working-sporting breeds from their greater populations for the specific purpose of public display and sales to the household market.

This is an important concept to understand. The modern hobby breeder specializes in a breed. A breed is a population of dogs that is mechanically isolated from all other dogs. Only those dogs registered in the breed stud book can be official members of the breed.

When I contrast this policy and process with those of working-dog breeders, who created the breed in the first place, I am horrified at what the hobby breeder has achieved. Breeders of working dogs did not attend dog shows to have their dogs judged on how they looked. They selected the best performers to breed dogs that were good workers. Most of them were not interested in "perfecting" a breed. As a sled dog driver I wanted to win races. What the dog looked like was important only for how that shape could run. Ancestry was important only for what it could indicate about a dog's potential to run.

Actually, of course, I did care how they looked. I wanted them to look like a Coppinger dog. It is a feather in the cap of any breeder of working dogs if his dogs develop a characteristic appearance, along with superior ability, which distinguishes them as being from a particular kennel. Breeds often are named for their breeder, for instance, the Reverend Jack Russell, or Ludwig Dobermann. Hound breeders, especially, strive for distinctive packs. Occasionally, and usually accidentally, a pup with a novel marker appears. A breeder might take a fancy to it and preserve it. The history of the golden retriever, as related in Chapter 3, illustrates this principle very well.

It also points up a very different process than the one producing pet or show dogs. A dog purchased from inbred stock (closed stud book), untested in the field for many generations, is the product of a breeding program (maybe) that has little to do with its working behavior. The expectation of the new owner is that the dog will be good because it is a purebred golden retriever. "What kind of dog shall I get?" "Get a golden retriever because they have a 'friendly nature and disposition, athletic ability, love of water, and natural instinct for hunting and retrieving.'"

What?! That sounds ridiculous to a working-dog person, or to a population geneticist. Friendly disposition is genetic? Love of water is genetic? Athletic ability has something to do with golden color? Is the implication that all goldens have this same set of genes, and all these traits? Is there no variation in golden retrievers? Lord Tweedmouth had good dogs because he had a good breeding program that included a high percentage of crossbreeding and because he hired people to work those dogs from their youngest days and develop the best dogs. He liked to hunt, he liked to have the best hunting dogs, and he was proud of his eye for working dogs. And he culled the bad ones. Anybody who ever created a breed did so by culling the ones they didn't want.

Today's household golden retriever is a caricature of Lord Tweedmouth's dogs.

The idea that there is something intrinsically desirable in the members of a breed is false. It is the same faulty notion as thinking there is something superior about "royal" blood. It is not only false, but it is bad genetics. It is not only bad genetics, but it dooms any breed that gets caught in that physically isolating trap.

The advantage for pet owners is that in a few generations of selecting for specific size, color, or other superficial reminders of the ancestral working dogs, any of the innate predispositions for the ancestral work almost certainly deteriorate. The rules of genetics say any character that isn't continuously selected for (say, working behavior) will begin to drift because of random events — disease, founder's principles, nongenetic calamities. In fact, what happens in the household world is that owners will quickly start selecting against many of the innate dispositions of a breed because of their obnoxiousness. The innate behaviors will begin to be described as compulsive behavioral disorders. Thus, dogs selected for their superficial traits might eventually make better household dogs because they lose their working dispositions.

But, however, and nevertheless, there is one other genetic mechanism operating. Closing the stud book on a population, in order to promote specific traits, inadvertently and dangerously starts a process of inbreeding. Inbreeding decreases the amount of genetic diversity. Genetic diversity is a source of genetic vitality. This genetic (phenotypic) vitality is good genetic health shown by an animal due to normal actions and interactions of the genes, uncomplicated by any deleterious effects that can be expressed when mutant gene products are paired. The chances of deleterious pairings are increased in inbred animals.

The first rule for all working dogs is they have to have stamina. Stamina is one result of genetic vitality. But stamina plus working behavior make for obnoxious pets. It is rare that any household can tolerate for very long an obsessive dog with stamina. Such a dog is usually much too active to be comfortable in the house. I am certainly not advocating this mechanism for selecting pets, but simply pointing out that sexual isolation and the inbreeding system of producing household dogs are certain over time to produce lethargic, i.e., peaceful dogs. Their personalities are amenable to the needs of their owners, even if their genotypes are in trouble.

There is an even more severe problem occurring in the genetics of the household dog. The working dog was selected to behave in a certain way. The sled dog, for example, was selected to run fast in harness with other dogs. In being selected for that behavior, the dogs evolved a unique shape. This shape allows the dog to behave fast, with stamina. The relationship between shape and behavior is omnipresent.

Therefore, if we want to change the behavior of a dog — make it more peaceful and less vital — we must also change its shape. Herein lies the dilemma for the breeder. The audience wants household dogs that are a historical representation of the working-breed shapes, and at the same time they do not want them to display working-breed behaviors. Trying to select for an acceptable household behavior while holding the working shape constant cannot be done. The dog will come apart. It will show genetic diseases. Its hips won't fit together right. The joints will show weaknesses, and the dog will twitch and bleed and each generation will become increasingly miserable.

Belyaev selected for tame foxes (Chapter 1), i.e., foxes with tame household personalities. And his foxes came apart. He couldn't select for tame and hold the wild-fox shape.

That is exactly what is happening to our household dogs.

Increasingly, the modern household dog becomes a genetic prisoner trapped in an isolated population. With each succeeding generation the behavioral and physical misfits get eliminated from the gene pool while breeders try to hold on to the ancestral form. But in each new generation we see a host of new genetic problems. Lists of breed-specific genetic diseases are now part of the professional and popular literature.

And it is worse than that. Breeders and owners forget what the historical dog looked like. They select for the exaggerated form. They select for the really big ones. They select for the flattest face. They select for the longest face. The breeds end up with weird conformations. Each breed takes on an unnatural shape, becoming a freak of nature. They are loved the way the hunchback Quasimodo was loved — a dichotomy between the grotesque form and the honorable personality.

As the decades go by, every part of the household dog's life is increasingly manipulated for the human host's benefit. The dog is capriciously manipulated for human pleasure. The more bizarre and exaggerated the animal is, the more benefit it seems to confer. This recent breeding fad for the purebred dog is badly out of control. It appears that selection for the exotic is the goal, probably to increase interest and sales. We are producing unhealthy freaks to satisfy human whims. This is terribly unfair to dogs.

The bulldog is perhaps the epitome of what I find alarming and unethical. Take the feisty and useful butchers' dogs of the seventeenth and eighteenth centuries, and turn them into sporting dogs. By the early eighteenth century, determine that tormenting a bull and risking the dog's life just for sport are unethical. The dog then is adopted as a pet-class dog. Over time each of the traits that made it a good cattle-catch dog for the neighborhood butcher gets blown so far out of proportion that the dog now resembles something out of a horror movie.

Their faces are so deformed they can't get their teeth to line up to chew. With some of the miniaturized forms there is no room for their teeth and they fall out upon erupting from the gums. Their faces are so squashed that the turbinate bones in their nostrils are tiny. Turbinate bones are covered with respiratory epithelial tissue, which helps the dog to breathe and cools its brain. As a result of the tiny turbinates, bulldogs and the other flat-facers have poor brain cooling, poor breathing, and low oxygen tension in their blood. Of the bulldogs that have been tested by the Orthopedic Foundation of America, 70 percent have hip dysplasia. None (zero percent) was reported with excellent hips.

Show-quality bulldogs often can't deliver puppies naturally. In fact, the dogs can't even breed, and have to be artificially inseminated. All of which causes pain. I once asked a woman at a dog show what her Boston terrier was good for. Without hesitation she said, "He is really good at snoring."

Obviously the original bull-baiting dogs didn't have any of these physical characters. And yet someone is always willing to tell the story that the modern bulldog has the turned-up nose so that it might breathe while clinging to the bull's neck. It is fiction. It is rationalizing the purposeful selection for the bizarre. But at the same time I hear people brag how their bulldogs can't even mate anymore. They are amused by these oddities.

The rationalizations for developing freaks are often as bizarre as the traits that result. Breeders propound far-fetched factoids: The komondor has hair hanging down over its eyes so that it can withstand the ultraviolet light bouncing off the snow on winter pastures; shepherds want that forty pounds of corded coat on their komondors to protect against wolf attacks; the great folds of skin on a shar-pei are to prevent serious injury to the underlying muscle in a fight; achondroplasia in dachshunds helps them go into holes after otters or minks; the great bulk of the Saint Bernard is needed to adapt them to tracking lost or hurt people in the Alps.

Of course, nothing could be further from the truth. These are made-up stories. Komondors and the other livestock-guarding dogs rarely have to fight wolves, and borzois and Scottish collies have such weak jaws and ill-formed mouths that they can hardly suck when they are born. There are lots of ways to get dogs into holes other than make achondroplastic dwarfs out of them.

All these traits are genetic deformations found to occur infrequently in all populations. The same genes for achondroplasia found in dachshunds and basset hounds are found in human beings. When the deformation shows up in people, nobody would have the audacity to say these people were selected to go into holes and kill badgers.

People who spend their lives working or hunting with dogs want dogs that look like dogs. There is a basic plan to dogs, and any dog that doesn't have that basic plan shouldn't be bred. The first statement out of any working-dog person's mouth is he wants stamina in his dogs. Certainly a dog with low oxygen tension, or fifty pounds of saggy coat, isn't going to show stamina. I certainly don't want my livestock-guarding dogs with hair hanging all over their eyes so they can't see. Neither do my sled dogs need blue eyes in order to run fast.

It is a terrible joke. Humans take total control of every aspect of the household-dog's life. They are bred capriciously to any shape, size, or color humans think is interesting or aesthetic. Their looks are conversation pieces. A rare breed takes on value for the sake of its novelty. Hobby breeders select for the visual impact of a dog in the show ring. That means amplifying or embellishing an existing trait. Magnify the size, the coat, the nose. Preserve the extra dewclaw, which supposedly helps a Pyrenean mountain dog stay on top of the deep snow in the mountains, but which I always clipped off my day-old husky pups so that as adults they would not tear their skin when they break through a snow crust. I once wrote a book where I designed dog freaks, like flounder hounders, a breed with both eyes on the same side of its head, and then I invented a job for this monster helping fishermen spot fish underwater. If I could figure out how to bioengineer that dog, I could make a lot of money.

A show champion in the household trade is not required to do anything except not bite the judge. Dog shows are comparable to human beauty-queen pageants. Compare each individual with the others in the show and see which one comes closest to some arbitrarily designated, idealistically "perfect" form. At least in the present human version of beauty contests, contestants have to say a few words, or play the clarinet, or recite a poem — to show a behavioral skill.

The difference between these competitions is that the human show has a minor effect on the population's genetics. The dog-show winner, however, becomes the favored breeding dog, and the tiny population that qualifies for inclusion in the breed's stud book is now funneled through these few "best" individuals. To breed heavily to champions is to substantially reduce the effective population size. It channels the available genes through a few individuals. If every owner of a female only bred his dog to this year's champion, then the next generation of dogs would all have the same father. Then the generation after that would all be brother-sister crosses. Thus, the breeding to champions increases the inbreeding coefficient very rapidly. On the surface some popular breeds may appear to have a large population, but their genealogies — their pedigrees — indicate that most of them share close ancestors and therefore also genetic alleles.

The household-dog world is not the only guilty party here. The same problems are now evident in the working-dog world. In the late 1970s I was buying border collies in Scotland, and looked far and wide to find a dog that did not have the champion herder Wiston Cap in its genealogy. (The dogs I found, by the way, were my pseudocompanion, Jane, and her brother Will.) Trialing border collies is a major sport in Scotland, complete with prime-time television coverage of the top trials, and also in the United States, with often generous prize money for the winners. Contestants are eager to succeed, and therefore seek to breed their dogs with the trial winners. Dogs from famous handlers and dogs that win command premium prices for stud services and puppies. But the genetic problems are increasing.

The same reduction in gene diversity takes place when a breed club tries to select against hip dysplasia, retinal atrophy, or some other so-called genetic disease. Every time an animal is culled for a genetic problem, the genetic variation in the closed population is further reduced. It's not just the bad genes that are affected, it is all the animal's genes. Any time there is selection for or against single characters, i.e., "tame" or "hip dysplasia," then one must be prepared for the appearance of new or altered characters because of what Darwin called "the mysterious laws of correlation." Today the phenomenon is called pleiotropy, or saltation — the fact that more than one characteristic can be controlled by a single gene, and selection can result in unintended and unpredictable changes.

When I look at the benefits for the dog in this symbiotic relationship with humans, it looks well-nigh hopeless. Many breeds are living to pay a terrible price for the temporal increase in population or the luxury of expensive food and care. It is not simply that the dogs have access to the kind of medical care that is given to humans, but that they have been bred so they need such care to survive. Breeds like the bulldog are in a dead-end trap. There probably is not enough variation left to get them out of their genetic pickle. Unless the breed clubs open their stud books and allow outside breedings, bulldogs and the other breeds caught in these eugenic breeding practices are headed for extinction. The problem here is that unlike the wild counterpart becoming extinct because of habitat loss, these purebred individuals will increasingly suffer ill health.

What is troublesome is that modern society seems to have little realization of what it is doing to dogs. Owners don't seem to be disturbed about deformation, or even that their dogs are overweight. They are pleased that this is their third Great Dane in ten years and appear proud of the fact that they can cope with its short longevity, giant size, and structural problems. Many, many people love their dogs right into obesity, having no idea of the discomfort from excess heat load caused by the fat.

I believe the modern household dog is bred to satisfy human psychological needs, with little or no consideration of the consequences for the dog. These dogs fill the court-jester model of pet ownership.

From the behavioral ecologist's point of view, I don't know what to call this symbiotic relationship. It may be a new category called reciprocal amensalism. It is bad for humans because of the economic and health problems created by large populations of dogs, and at the same time it is bad for dogs. Individual people may get some psychological benefit, and I suppose there is always a chance for some dog to have a life of true luxury. Nevertheless, for purebred dogs the scope of their existence is to be chosen (or not) according to principles of eugenics — of the worst kind. The breeding programs are not concerned with adapting the dog to the household environment. Rather, the dog is being bred for its showplace value, a not-so-mere-bagatelle of form, with little concern for what's inside, or even if the animal inside that aesthetic shape hurts.

It's a bestial way to treat your best friend.

Copyright © 2001 by Raymond Coppinger and Lorna Coppinger

Table of Contents



Preface: The Right Kind of Dog

Introduction: Studying Dogs
Why Study Dogs?
How to Study, and Who Studies, Dogs

Part I. The Evolution of the Basic Dog: Commensalism

Chapter 1. Wolves Evolve into Dogs
The Pinocchio Hypothesis of Dog Origin
Taming the Wolf
Training the Wolf
Domesticating the Wolf
Speciation Requires Populations That Evolve — Not Individuals
Speciation Requires Differential Mortality

Chapter 2. Village Dogs
The Mesolithic Island

Chapter 3. Natural Breeds
People Become Conscious of Dogs

Part II. Working Dogs and People: Mutualism

Chapter 4. Developmental Environments
Livestock-Guarding Dogs
In the Nest: Shaping the Behavior
The Transhumance: Distributing and Mixing Genes
The Transhumance: Evolving the Size and Shape
Breed Genesis: Selecting for Color
Walking Hounds

Chapter 5. The Physical Conformation of a Breed
Sled Dogs — How Do They Run?
The Shape of the Team
Running Is Social Behavior
The Society of a Sled Dog Team
The Value of the Breed Standard

Chapter 6. Behavioral Conformation
Herding Dogs, Retrievers, and Pointers
The Border Collie's Behavioral Conformation
Motor Patterns

Part III. Are People the Dog's Best Friend?

Parasitism, Amensalism, and Dulosis

Chapter 7. Household Dogs
Measuring the Benefit to Humans of the Household Dog

Chapter 8. Assistance Dogs

Part IV. The Tail Wags the Dog

Chapter 9. What's in the Name Canis familiaris?

Chapter 10. The Age of the Dog

Chapter 11. Why Dogs Look the Way They Do
How to Change Size
How to Change Shape
The Shape of Intelligence
Rapid Evolution of Breeds
Neoteny, Paedomorphism, and the Evolution of Dogs from Wolves

Chapter 12. Conclusion



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