dog foot print and dna diagram

Essays on Dog Genetic Diversity

Inspired by conversations hosted by Jemima Harrison

The purpose of this website is to host essays and information about maintaining viable populations of healthy pure bred dogs

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copyright © 2009 by Dr Susan Thorpe-Vargus


Have you ever wondered at the extraordinary diversity in the appearance of various dog breeds? How is it that a Yorkshire Terrier can be the same species as a Bullmastiff, or a Pug be related to a Saluki? What are the factors that have led to this incredible range and variety in appearance, not to mention behavior and temperament? It is not simply a question of phenotype vs. genotype, or dominant vs. recessive genes. Let’s begin the journey by looking at how dogs evolved into the companion we know and love today.


About 60 million years ago a small weasel-like animal lived in many parts of Asia. This ancestor of all modern day canids (dogs, jackals, wolves and foxes) was called Miacis. Cynodictis, the first true dog-like canids are thought to have descended from Miacis about 30 million years ago. This line eventually split into two branches, one in Africa and the other in Eurasia. The Eurasian branch was called Tomarctus and was, until recently, thought to be the progenitor of wolves, dogs, and foxes. However, new research has called this theory into question with a recent paper indicating now that the wolf is the domestic dog’s only direct ancestor and that a recently shared ancestry with the fox and jackal is unlikely. 2 This somewhat controversial paper also asserts that the first domestication of wolves may have taken place as long as 100,000 years ago. The actual time that such domestication occurred, of course, cannot be settled based solely on mtDNA analysis. Indeed, the new data does not clearly support that the dog is descended from the wolf. Neither do the fossil remains. A case can still be made that they coevolved from a common ancestor.

Research now suggests that the domestic dog line began to diverge from the wolf after the first wolf became domesticated. Over time, groups of wolves became adapted to a niche that made them ultimately better suited to domestication at some point as early as 100,000 years ago to as late as 14,000 years ago. The actual timing remains in dispute since the fossil records are not consistent enough to pinpoint an exact period of time. However it has been well established now that different domestication events did occur from multiple populations by researchers such as Robert Wayne3. This makes sense as both wolves and humans coexisted over a wide geographical area and it is likely that multiple domestication opportunities would have arisen. These multiple events in various parts of the world accentuated the diversity we see in dogs today.

As hunter/gatherers, humans would have found dogs very useful. Then, about 8,000 years ago, humans turned to a more settled way of life. This is when severe selection for specific behaviors and traits became important and ‘modern’ breeding practices started. And so it begins – dogs bred for many reasons, from companionship to guarding abilities and so on.

The concept of a “pure” breed is a relatively recent one; further back, local dog populations consisted of similar looking dogs bred for a specific purpose. Although there were some exceptions, the dog breeders of that time did not hesitate to breed a dog of one type to a newly arrived dog from another area. Thus, up until the 19th century the various dog breeds were more often than not strains of closely related and similar looking dogs that as a population had a great deal of genetic diversity. Only dog populations that lived in geographic isolation approach today’s purebreds in terms of a restricted gene pool.

In searching for cultures without dogs since pre-historic times we come up empty handed. Thus we find early recognizable breeds coming from the Middle East, Africa and Asia. The Middle Eastern coursing hounds had become well established no later than 2,000 BC. The Basenji, a hunting breed of the African savannah, may predate the dogs of the Pharaohs. In the Far East, isolated areas such as Tibet and Mongolia produced a number of still extant breeds of ancient origin. Malta was occupied as early as 3,500 B.C and the dog brought to Malta may have had earlier Egyptian origins. The point here is that since relatively early times in recorded history, there has been a tremendous diversity in dogs. Contrast the Roman Mollosus – a mastiff-like creature (or what we think it looked like)--with the Maltese or the Tibetan terrier or the Lhaso Apso, and it immediately becomes obvious that there may be no “standard” dog. The tiny kingdom of Tibet, produced many different breeds, some now probably extinct, but which include the following breeds and their ancestors: Kuvasz (before Hungary); Lhaso Apso, Tibetan Terrier, Tibetan Spaniel, Tibetan Mastiff, just to mention a few familiar to Western dog fanciers. Even dogs we think of as “English” such as Mastiffs have had ancient origins. Recognizably mastiff-like dogs can be seen on Egyptian monuments circa 3,000 BC. They were in China circa 1100 BC and eventually went to England with invading Roman forces in the first century AD. We can safely say that certain dog strains have been breeding true for a very long time.

NEOTENY – More Fuel For Diversity

There is a saying among dog breeders that “All puppies look alike.” Newborn puppies of different breeds, except for size and of course color, look remarkably alike. How is it that they grow up to look so different from one another? The vast array of physical and behavioral differences in dogs is probably not due to selection for each individual trait, but more likely to selection for groups of traits that are all similarly affected by the same hereditary mechanisms. One such mechanism is the regulation and timing of developmental processes. Selection for one trait affected by developmental timing could inadvertently select for other traits also thus affected.

It is very likely that this process has played a vital role in the initial domestication and later diversification of dogs. As animals mature, they pass through different stages, each uniquely adapted to its particular circumstance. In wolves, neonates and juveniles are dependent upon parents to care for them, and they are extremely successful at eliciting that care. In comparison to adults, they are relatively tame and subservient. Wolves (or the wild ancestors of wolves and dogs) that tended to retain these immature qualities of tameness and subservience into adulthood would have been favored by early humans and would have formed the core of primitive domesticated dogs. This retention of immature characteristics in adults is known as neoteny. By choosing the individuals to reproduce that showed the favored immature behavioral qualities, concurrent selection for other juvenile traits -both wanted and unwanted--may very likely have occurred, laying the basis for the diversity seen in dogs today.

The rounded head and shortened muzzle of some breeds is reminiscent of the neonatal wolf. Floppy ears, too, are a neonatal wolf trait. The dog’s smaller head, brain, and teeth, in comparison to wolves, are comparable to those of the immature wolf. Many of the herding, hunting, and playing behaviors humans have found so useful and entertaining can be found in immature forms of hunting behavior in wolves. Barking is rarely seen in adult wolves, but is a trait of juveniles---as well as adult dogs. Further crossing of dogs showing differing degrees and influences of neoteny could produce novel combinations of adult and immature characteristics, so that domestic dogs may be regarded as a blend of immature and adult characteristics. Sometimes this creates problems for the dog breeder as in the case of toy breeds with disproportionately large eyes. The eye seems to be relatively immune to neoteny and is therefore difficult to reduce in size through selection in contrast to body and skull size, both of which have been induced to retain immature dimensions.

Now, let’s take a look at some of the basic concepts of Genetics that every breeder needs to know. Then we'll discuss the mechanics of inheritance in Chapters 2 and 3.

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Articles at

Blogs Discussion Controversies in Dog Breeding

Jemima Harrison

Pedigree Dogs Exposed - The Blog

Border Wars

A Border Collie Manifesto

Desert Windhounds

A Bitter Cynoanarchist Rages On

Carol Fowler

Dog Breed Health

Interactive Genetics Tools Websites

Online COI Generator by Henk Meijers

Online COI Calculator

Academic Genetics Tutorials/Courses/Resources Online

Online Mendelian Inheritance in Animals (OMIA) OMIA

Understanding Gene Loss in successive generations An Interactive Academic Site on "Gene Dropping" at University of Montana

BIOL 4312: Genetics

GENUPComputer aided learning for quantitative genetics.
Although a Windows application Bonnie Dalzell has been able to run it under Ubuntu Linux from my desktop.

Design Of Crossbreeding Programs

A tactical approach to the design of crossbreeding programs.

Applied Animal and Plant Breeding

Cultivar Crazy - Preserving the Genetic Heritage of Plants

Applied Animal and Plant Breeding

Fitness and Viability of Small Populations
by Nina Pekkala

The Effects of Genetic Drift, Inbreeding, and Interpopulation Hybridization experimental study involving Drosophila (fruit flies)

Authoritative Inherited Diseases in Dogs Sites

Inherited Diseases in Dogs

Inherited Diseases in Dogs (IDID)

Canine Inherited Disorders Database

Canine Inherited Disorders Database

Inherited Disorders in Animals

List of Inherited Disorders in Animals (LIDA)

Canine Health Registries and Organizations

VMDB Veterinary Medical Database
The Veterinary Medical Database (VMDB) compiles patient encounter data from nearly all North American veterinary medical databases.

CERFCanine Eye Registration Foundation

CHIC) Canine Health Information Center

OFAOrthopedic Foundation for Animals

CERF Canine Eye Registration Foundation

APGAW: Associate Parliamentary Group For Animal Welfare
[Great Britain]

A Healthier Future for Pedigree Dogs

Canine Genetics Sites

The Hounds of Claybrook

Closed Registries, Genetics, and Inbreeding Depression


Canine Genetics Resources

Canine Diversity Website Homepage

Canine Diversity Homepage

A series of articles concerning the genetics of selection, inbreeding, inheritance of congenital defects.

Canine Inherited Disorders Database

CID Database

Sue Ann Bowling:

Animal Genetics Pages

To Breed or Not to Breed

All 4 essays by Susan Thorpe-Vargas Ph.D., John Cargill MA, MBA, MS, and D. Caroline Coile, Ph.D

The Genetic Cul-De-Sac

Dogs As An Endangered Species

Breed Related Sites

Borzoi Heritate Borzoi Pedigree Collection

Borzoi Pedigrees

Cesky Terriers

Cesky Terriers

The Dalmation Heritage Project

Project to breed Dalmations with normal uric acid metabolism

Author's Sites in no particular order:

John Burchard

Tepe Gawra Salukis

Jim Seltzer

Willowind Dalmations

Jeffery Bragg

Seppala Siberian Sleddogs

Dorothea Penizek

Clarmorris Parson Russell Terriers

Bonnie Dalzell

Borzoi Information Site

Academic Articles on Inbreeding, Fitness and Population Genetics

Fitness and Viability of Small Populations

by Nina Pekkala

The Effects of Genetic Drift, Inbreeding, and Interpopulation Hybridization experimental study involving Drosophila (fruit flies)