Canine Behavior: Insights and Answers 2nd Edition

319 Pages·2009·13.04 MB·English

by  Bonnie V. Beaver BS DVM MS DACVB

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CANINE BEHAVIOR: INSIGHTS AND ANSWERS, SECOND EDITION                     ISBN: 978-1-4160-5419-1
Copyright © 2009, 1999 By Saunders, an imprint of Elsevier Inc.
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Library of Congress Cataloging-in-Publication Data
Beaver, Bonnie V. G., 1944-
Canine behavior : insights and answers / Bonnie V. Beaver. -- 2nd ed. p. cm.
Includes bibliographical references and index.
ISBN 978-1-4160-5419-1 (pbk. : alk. paper)
1. Dogs--Behavior. I. Title.
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Veterinarians work to make the lives of animals and people better—it is our passion. We do it
as we heal the sick, ensure a safe food supply, protect our country, gather new knowledge, teach
the next generation, and tackle behavior problems. Our profession is small in numbers but
mighty in accomplishments.
Since the last edition of the book, I had the privilege of serving my profession as the Chair of
the Executive Board and then President of the American Veterinary Medical Association. It is
a long way for a little girl from rural Minnesota to come, especially in an era when she was told
that “women cannot be veterinarians.” It was an unbelievable journey for which I am truly
grateful. I dedicate this book to all my veterinary colleagues. Without your support I could not
have accomplished this book nor had my AVMA leadership experiences.
Preface
Considering the many recent advances in the un-  To understand abnormal, it is important to 
derstanding and application of behavioral science  have a clear understanding of normal. The chapters 
and modification principles, a revision of Canine are organized around various large categories of be-
Behavior is definitely overdue. This second edition  havior. Within each chapter, normal behaviors are 
is intended to explore the many facets and conse- described first, followed by the various types of re-
quences of behavior in dogs—from normal to ab- lated behavior problems. Because dogs live in a wide 
normal—as it has developed over time. Even though  variety of environments, it is impossible to include 
normal behavior does not essentially change, sig- every scenario of behavior that might occur, but 
nificantly new information is being gathered about  several that have been identified over the years are 
our awareness of the complex behaviors of the dog,  included. This edition discusses many of the classic 
starting with the dog genome and genetic changes  normal and abnormal behaviors and expands upon 
from a wolf-like ancestor to the many varieties of  current understandings of major problems such as 
dogs we recognize today. There have also been ma- stereotypies, cognitive dysfunction, and separation 
jor strides in knowledge about how to treat a wide  anxiety. It also presents the latest insights and ap-
range of problem behaviors, which reflect our con- plications of behavioral pharmacotherapy as well 
stantly changing relationships with these magnifi- as practical, rational, and humane approaches to 
cent creatures and are specifically addressed in this  modification techniques for basic and complicated 
edition. behavioral problems. 
It is my aim to meet several goals with this edi- Behavior problems result in the death of approx-
tion. Firstly, the content represents critical and  imately 10% of the dog and cat population each 
valuable material gleaned from a vast body of clas- year—that’s more than caused by all infectious dis-
sic and newer scientific references. This is intended  eases. As dogs have changed their habitat from the 
to help researchers, teachers, veterinarians, and pet  barnyard to the backyard and into the house, the 
owners gain further insights into the dog’s commu- importance of their behaviors and consequences 
nicative, social, sexual, ingestive, eliminative, loco- has been amplified in the lives of their owners.
motor, grooming, and sensory behaviors. Secondly,  Veterinarians  must  be  able  to  advise  clients 
it is designed to provide practical information to  about normal behavior and help prevent and eradi-
veterinarians for the diagnosis and treatments of  cate or minimize problems associated with abnor-
various behavioral problems. This is accomplished  mal behaviors. Only by knowing the natural history 
by providing detailed insights into normal dog be- of the dog and its innate behavior influenced by the 
havior, how problem behaviors develop, and how  modern realities of domestication can we serve the 
they might be prevented and managed successfully  needs of our canine patients so that they remain 
and humanely. Lastly, but not least, I hope the book  valued members of our households and communi-
will provide interesting information for fellow dog  ties for a long time.
enthusiasts.  Bonnie V. Beaver, DVM, MS, Dipl. ACVB
Acknowledgments
Many people have helped, both directly or indirectly, to ensure the publication of this book. 
Their support and backing have been important.
My appreciation is expressed to all of my colleagues, particularly Richard Adams, Sandee 
Hartsfield, Deb Zoran, Nini Hodges, Lore Haug, Patty Hug, and M.A. Crist.  Special thanks 
go to Peppy Shortstockings, Murphy Brown, Rachael Brown, Thelma Beaver, Shirley and 
Tommy Clem, Larry and Sherry Piper, Lauren and Stan Stephenson, Jean and Nathan Pip-
er, the John Smith family, George and Nyla Rayburn, and the AVMA staff.  Finally, I wish 
to thank the staff at Elsevier for their patience, particularly Maureen Slaten for her ready 
assistance and her gentle but persistent reminders of important deadlines.  I did make it.
Chapter 1
Introduction to Canine
Behavior
Understanding the dog requires that one under- larger brains.60 the line of ancestors that followed is
stand where it came from, so it becomes necessary less clear. Some propose that the Cynodesmus of the
to go back to the wolf and beyond. the dog began Miocene gave rise to the Tomarctus of the pliocene,
its association with humans well over 10,000 years and the Tomarctus eventually gave origin to the ca-
ago, a period longer than for any other domestic nids.60,89 Others theorize that Hesperocyon gave rise
animal. Since dogs came into our lives, they have to the Leptocyn, one possible ancestor of the dog, or
filled a number of roles: from companion to food, to the Tomarctus, another possible ancestor.182,211
from alter ego to special sense. It is the great genetic regardless of the exact ancient line, all mod-
plasticity in Canis familiaris that has permitted so ern predators (fissipeds) belong to one of three
much variation in its sizes, colors, and behaviors. In superfamilies: Miacoidea, Feloidea (aeluroidea),
modern times, breeds have been developed for spe- or Canoidea (arctoidea). Cats, hyenas, and Old
cific purposes, such as herding cattle, retrieving a World civets belong to the Feloidea. Dogs, wolves,
hunter’s kill, and pulling a sled. although there are foxes, and jackals belong to the canine subfamily of
behavioral variations between breeds, often related Canoidea.60,78
to purpose, the core behaviors of the domestic dog even narrowing down the phylogenetic subfam-
are very similar across breeds and are often simi- ily does not settle another age-old question. What
lar to those of its closest relative, the wolf. the wolf animal is the immediate ancestor of the domesti-
can serve as a basis for understanding the dog, but cated dog? Most experts agree that the wolf played
it is also important to remember that thousands of a major role in the dog’s genetic pool, but there are
years of selective breeding have changed a lot of the 32 subspecies of Canis lupus.211 Of the canids, only
behavior too. the wolf, coyote, and jackal have the same number
of chromosomes (78) as the dog and can interbreed
to produce fertile offspring.57 a question remains
HISTORY OF CANINE
about other common early ancestors or another
DEVELOPMENT
possible ancestor species.89,90,318 Dogs and wolves
the oldest known ancestor of the domestic dog share 71 of 90 behavioral patterns, more than are
is the Miacis, a small, weasel-sized carnivore that shared by any other two species.279 early DNa tests
lived approximately 40 million years ago in the have provided limited information that the domes-
eocene.60,88,89,211,239 the miacid was probably a tic dog, wolf, coyote, and jackal are the most alike
forest dweller with a long tail, short limbs, and a among the canids.69,317,326 Newer isoenzyme ge-
plantigrade stance.60,210 already at this time, the netic distance indicates that the dog is more closely
teeth that would come to be called the carnassial related to the wolf than to other canids.119,183,314,323
teeth had taken on their characteristic shape. By the assuming the wolf (Canis lupus) begot Canis lu­
late eocene period, the Cynidictus had evolved, and pus familiaris,211 which begot Canis familiaris, which
it probably gave rise to the Hesperocyon (formerly of the many subspecies of wolf started the process?
Pseudocynonsides).60,89,182,211 this latest group again, considerable disagreement exists. Some
existed approximately 3 million years ago in the authorities argue for a single origin in northern
Oligocene and was distinguished by its larger size, eurasia,278 the Near east,69 or North america.119,211
longer limbs, better developed carnassial teeth, and Others argue for the possibility of the occurrence

 Chapter 1 Introduction to Canine Behavior
of domestication in different species in diverse exactly when humans started interacting with
locations.55,57,85,190 how the first dogs gradually wolves is difficult to determine. Some of the ear-
changed into the many breeds we have today has liest hominid sites with Canis lupus remains were
been the subject of much speculation over the found in Zhoukoudian, China, dating back be-
years. Most people believe that the arctic breeds tween 200,000 and 500,000 years; in Kent, england,
probably came first because their physical appear- dating back 400,000 years; and in Nice, France, dat-
ances are very similar to wolves. Others believe that ing back 125,000 years.59,211 as evidence of domes-
“village dogs” were first.35,65 these were followed tication, scientists have pointed to (1) the smaller
by livestock-guarding dogs and hunting compan- size of bony remains compared with those of in-
ions, and then by sled dogs, herding and gun dogs, digenous wolf populations; (2) certain anatomic
and household dogs.65 the most current studies differences in the skull, mandible, and teeth; and
of DNa microsatellite subtypes find at least four (3) phylogeographic variation. according to these
distinct groupings representing unique “adaptive criteria, dogs separated from wolves up to 40,000
radiations” (Fig. 1-1).230 the four groups with sim- years ago. however, DNa evidence for the delin-
ilar genetic clusters also tended to share geographic eation suggests that dogs originated approximately
origin, morphology, and/or genetic closeness to the 135,000 years ago and that occasional wolf back-
wolf of Southeast asia, following humans to differ- crosses have occurred.313
ent parts of the world.35,230,267 research into com- Domestication probably occurred in more re-
mon behaviors also shows some similarities within cent history, but exactly when and where is not
the four groups. those closest to wolves show more clear. Burial sites are the most likely source of in-
aggression and less demand for affection than those formation, but some researchers question if burials
farthest away.111 truly indicate early domestication, or if they were
Hoary Fox
Crab-Eating Fox
Maned Wolf
Canoidea
Bush Dog
Black-Backed Jackal
Cape Hunting Dog
Coyote
Gray Wolf
Chinese Shar-Pei
Shiba Inu
Chow Chow
Akita
Basenji
Siberian Husky
Alaskan Malamute
Afghan Hound
Saluki
Mastiff type
All other breeds Herding type
Recent European origin breeds
Figure 1-1. Genetic subdivisions of Canids. (Data from references 230, 326, 327.)
Chapter 1 Introduction to Canine Behavior 
started somewhat later. a German dog burial site favored status and were more apt to be around oth-
dates back 14,000 years.58,93,238 another burial site ers with similar traits.
in Siberia goes back 10,650 years.93 evidence of the early humans may actually have taken young
dog’s existence has been found at the palegawra cubs from the den to be hand-raised, as was done
Cave in Iraq, dating back some 10,000 to 12,000 by Native americans even into the eighteenth cen-
years,55,56,211,212 although there is some question tury, and as some people still do.56,100,182,280 With-
about this date.69 at ein Mallaha in northern Israel, out a parent to teach hunting skills, the wolf/dog
scientists excavated a tomb dating back to 9350 would have become dependent on the humans for
to 9750 bc that contained a burial of an old hu- food.40,190 Some probably became scavengers, re-
man, probably a woman, and a 3- to 5-month-old sponsible for keeping the camps clean.57,90 as sug-
puppy.57,59,69,71,190 In China, domesticated dogs gested by evidence from the Israeli burial site, some
have been dated back to 7355 ± 100 bc.211 the wolves may have first shared space with humans
earliest european site is Star Carr in Yorkshire, simply to provide companionship.57,69 the relative
england, which contained evidence of dogs dating lack of canid bones at most sites indicates that dogs
back to 7538 ± 350 bc.69,211 Swedish and Dan- were probably not a major food source.69 Other
ish sites date back to 5000 to 6800 bc.211 In North roles these animals could have filled would have
america, dogs probably accompanied humans over been as an occasional “watchdog,” religious symbol
the Bering Strait land bridge from asia and spread or sacrifice, draft animal, or source of hides, meat,
out from there.165,230,238 Dog burial sites date back or warmth at night.57,90,182,190
8500 years in western Illinois and 7500 years in Ben- Domestication is a complicated process that in-
ton County, Missouri.69,199,210 the ancient Chirib- volves genetic changes in a large number of animals
aya dogs of peru were apparently very prized and that are selectively bred over many generations to
recently found mummified bodies date back about intensify certain traits and make others secondary.
1000 years.61 Domesticated dogs probably did not the result is a species that is biologically changed in
exist in australia more than 12,000 years ago.57,69 its morphology, physiology, and behavior.94,157 the
that was the approximate time tasmania separated dog became smaller than the wolf, had different
from the australian mainland, and the absence of colored hair, and was less alert to its environment.
canids there supports this theory. the dogs that did Over time the quantitative nature of the various re-
come to the australian continent probably did so as sponses change so that rather than a behavioral trait
recently as 5000 years ago,70 and eventually reverted being completely lost, its threshold is heightened or
to the feral dingo. lowered by lack of or consistent exposure to specific
how and why domestication of wolves occurred stimuli.249 Many more generations of continued se-
will always be speculative. Some theories about lective breeding modified the dog into a wide vari-
domestication in general suggest that a mutually ety of shapes, sizes, colors, and behaviors based on
advantageous relationship between wolves and hu- the goals and desires of those responsible for mate
mans would allow for the loss of such wolf behav- selection.
iors as self defense and self protection. at the same the domestication process can occur relatively
time, there would be the gain of other behaviors, rapidly in a canid when strict criteria are used.249
such as access to food or shelter.40 perhaps wolves the russian geneticist D.K. Belvaev was able to cre-
and humans shared hunting success. Wolves might ate major changes in a group of silver foxes through
have followed hunters initially to find game. the intense selective breeding only for tameness over
wolf that was allowed to share a kill made by its approximately 20 generations.26 By the end of that
two-legged benefactor would have become some- time, his animals were reported to have sought
what tame.211 hunting skills as well as ease of being contact with humans, licked human hands and
tamed—traits that benefited the humans—could faces, changed their breeding cycle to twice a year,
then have been selectively bred owing to proxim- developed drooping ears and erect tails, and dis-
ity because the most successful wolves took on a played the submissive behaviors of whining and tail
 Chapter 1 Introduction to Canine Behavior
wagging.182,190 In the case of wolf-to-dog evolution, and are able to read human signals, including those
the timeframe was probably much longer than in that indicate the location of hidden food.105 even
Belvaev’s experiment, because the selection criteria wolves raised with humans do not show that be-
were far less distinct; however, the social structure havior. Coat colors have increased in dogs. It has
of the wolf is the closest of any of the wild canids to been shown in foxes that the change in the color of
that of the human, making the wolf the best candi- fur can be related to behavior changes. the amber
date for domestication.278 fox is less aggressive than the gray fox. also, at least
the physical and behavioral features that tended some amber foxes have smaller adrenal and pitu-
to be retained in the wolf-to-dog transition were itary glands.57
those most associated with juveniles, a phenom- Seven thousand years ago, there were specific
enon called paedomorphosis.* In the dog, those kinds of dogs. three thousand years ago heavy hunt-
wolf cub–like physical features include a smaller ing dogs, short-legged dogs, and Greyhounds ex-
size, shorter muzzle, relatively wider cranium or isted.57,58 Soon dog breeds proliferated. Small dogs
more domed head, reduced tympanic bullae, more share a single insulin-like growth factor I (IGF-I)
convex mandible, larger eyes, smaller teeth, floppy allele that suggests a specific sequence variant is the
instead of erect ears, and less developed tempora- major reason for their body size.298 But dogs now
lis and masseter muscles. † the retained behav- come in a wide variety of sizes, colors, and working
ioral characteristics include increased frequency in skills. Dogs were an integral part of roman culture
care-soliciting behaviors, submissive food begging and could be found throughout that empire. prob-
(such as face licking), relative lack of fear, tim- ably the first systematic breeding of dogs occurred
ing of activity periods, curiosity, playfulness, less- there. the romans had fighting dogs, sheepdogs,
ened territorialness, and increased seeking of social guard dogs, and small lapdogs.56,57 as a side note,
contacts.40,87,119 the cave canem signs in pompeii and rome were
On the other hand, some distinct differences the first “Beware of the Dog” signs; their intent
also developed between wolves and dogs that were was to warn visitors not to step on the small Italian
unrelated to paedomorphosis. Many of these fea- Greyhounds.239 Columella, a roman agricultural
tures were not selected for or against but probably authority, is said to have advised the breeding
tagged along genetically. the curly tail of some dogs of dogs that did not look like wolves, including
was one of the most obvious early changes. It, in black farm dogs that barked at intruders and white
turn, has been followed by a great variation in the sheepdogs.56
size and shape of the tail, its amount of curl, and the licensing of dogs was started by King henry
the position in which it is carried. Unlike other III of england around the middle of the thirteenth
canids, dogs sometimes have the first digit on the century when he singled out mastiffs.239 the first
rear limbs.57 the scent gland from the dorsal aspect competitive dog show was held in 1859 at Newcastle
of the base of the tail is greatly reduced or absent in in Britain but was limited to pointers and Setters.
dogs.57,94 reproductively there are differences too. With the start of the shows and the British Kennel
Sexual maturity occurs between 6 and 12 months of Club in 1873, breed standards were gradually tight-
age in dogs versus almost 2 years in wolves. Female ened until they reached what we know today.57
dogs experience twice as many estrous periods as
female wolves in a year, and estrus can occur at any
CURRENT STATUS OF CANIDS
time during the year. thus, males are reproductively
interested throughout the year instead of during a Dogs have been used to meet more human needs
wolf-like 2-month breeding season.57,278 Dogs have than any other domestic species (table 1-1), and
developed better social-cognitive skills than wolves humans keep finding new things for them to do.
the original list of four roles animals typically fill—
food, clothing, shelter, and transportation—were to
*References 40, 57, 87, 94, 190, 249.
†References 40, 56, 57, 59, 94, 190. be greatly expanded as dogs came to be associated
Chapter 1 Introduction to Canine Behavior 
TABLE 1-1    Primary roles played by domestic animals throughout history
Primary roles Dogs Cats Horses Cattle Swine Sheep Goats Camels Reindeer
Clothing/shelter
Fiber + + + +
Hides + + + + + + +
Communication +
Draft or power + + + ± +
source
Educator + + + + + + + + +
Extension of + + +
human senses
Food
Meat + + + + + + +
Milk + + +
Blood + +
Manufacturing
Feces for energy + + + +
Feces for feed + +
Health products + + + +
Occupation + + + + + + + + +
Physical health
Poison detection +
Exercise + + +
Protection +
Psychological
health
Companionship + + + + + + + + +
Extension of ego + + + + + + + + +
Recreation + + + + + + + + +
Religious figure + + +
Therapy + + + +
Research + + + + + + +
Transportation + + + +
with other cultures and time periods. a recognition integration. their use has been correlated with bet-
of the human-animal bond allows for consideration ter attendance at work or school, while reducing
of activities animals do with us, as well as for us. prob- the number of human assistance hours needed.5
ably the longest service that dogs have performed is Dogs are also an integral part of many search-and-
the guiding and guarding of livestock. even today rescue teams.118 they have served with honor in the
the use of livestock-guarding dogs is an effective military as sentries, smell detectors, early warning
method of managing predation, with 82% of them systems, and communication go-betweens.4 Dogs
considered to be an economic asset.103 Guide dogs are used as research models and as a child’s teacher.
for the blind are the most common type of specially psychologically, they can contribute to good men-
trained service dog, extending the sensory capacity tal health, help bridge social interactions5,168 and
of the human with whom they work. But service serve as a warm, fuzzy therapist for those who need
dogs help humans in much more than the physical a feeling heart to listen.
sense. they have been shown to provide a tremen- the genomics of dogs is currently a hot re-
dous psychologic boost and to facilitate community search topic, but ultimately the understanding of