Origin of the domestic dog

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The origin of the domestic dog (Canis lupus familiaris and sometimes referred to as Canis familiaris) began with the domestication of the wolf tens of thousands of years ago.[1][2][3] Genetic and archaeological evidence shows that humans domesticated wolves on more than one occasion, with the present lineage of C. l. familiaris arising at the latest 15,000 years ago as evidenced by the Bonn-Oberkassel site and possibly as early as 33,000 years ago as evidenced by the mtDNA testing on a paleolithic dog's remains from the Razboinichya Cave (Altai Mountains).[4][5][6][7][8][9][10] Some research indicates that dogs may have diverged over 100,000 years ago.[11]

Domesticated dogs provided early humans with a guard animal, a source of food, fur, and a beast of burden; humans provided the dogs with food and shelter. The process continues to this day, with the intentional artificial selection and cross-breeding of dogs to create new breeds of dogs.


The earliest fossil carnivores that can be linked with some certainty to canids (wolves, foxes and dogs) are the Eocene Miacids some 38 to 56 million years ago. From the miacids evolved the cat-like (Feloidea) and dog-like (Canoidea) carnivores. The canoid line led from the coyote-sized Mesocyon of the Oligocene (38 to 24 million years ago) to the fox-like Leptocyon and the wolf-like Tomarctus that wandered North America some 10 million years ago.[12]

Canis lepophagus, a small, narrow skulled North American canid of the Miocene era, led to the first true wolves at the end of the Blancan North American Stage such as Canis priscolatrans which evolved into Canis etruscus, then Canis mosbachensis,[13] and in turn C. mosbachensis evolved to become Canis lupus, the immediate precursor to the domestic dog.



How exactly the domestication of the wolf happened is unclear, but theories include the following:

Orphaned wolf-cubs

Studies have shown that some wolf pups taken at an early age and reared by humans are easily tamed and socialized.[14] At least one study has demonstrated that adult wolves can be successfully socialized.[15] However, according to other researchers attempts to socialize wolves after the pups reach 21 days of age are very time-consuming and seldom practical or reliable in achieving success.[16]

Many scientists believe that humans adopted orphaned wolf cubs and nursed them alongside human babies.[17][18] Once these early adoptees started breeding among themselves, a new generation of tame "wolf-like" domestic animals would result which would, over generations of time, become more dog-like.[19]

Promise of food/self-domestication

Early wolves would, as scavengers, be attracted to human field kills and refuse left at human campsites.[20] Dr. Raymond Coppinger of Hampshire College (Massachusetts) argues[21] that those wolves that were more successful at interacting with humans would pass these traits on to their offspring, eventually creating wolves with a greater propensity to be domesticated. The "most social and least fearful" wolves were the ones who were kept around the human living areas, helping to breed those traits that are still recognized in dogs today.[18]

Coppinger believes that a behavioral characteristic called "flight distance" was crucial to the transformation from wild wolf to the ancestors of the modern dog. It represents how close an animal will allow humans (or anything else it perceives as dangerous) to get before it runs away. Animals with shorter flight distances will linger, and feed, when humans are close by; this behavioral trait would have been passed on to successive generations, and amplified, creating animals that are increasingly more comfortable around humans. "My argument is that what domesticated—or tame—means is to be able to eat in the presence of human beings. That is the thing that wild wolves can't do."[22]

Furthermore, selection for domesticity had the side effect of selecting genetically related physical characteristics, and behavior such as barking. Hypothetically, wolves separated into two populations–the village-oriented scavengers and the packs of hunters. The next steps have not been defined, but selective pressure must have been present to sustain the divergence of these populations.

Experimental evidence

As an experiment in the domestication of wolves, the "farm fox" experiment of Russian scientist Dmitry Belyaev attempted to reenact how domestication may have occurred.[23] Researchers, working with wild silver foxes selectively bred over 35 generations and 40 years for the sole trait of friendliness to humans, created more dog-like animals. The "domestic elite" foxes are much more friendly to humans and actually seek human attention, but they also show new physical traits that parallel the selection for tameness, even though the physical traits were not originally selected for. They include spotted or black-and-white coats, floppy ears, tails that curl over their backs, the barking vocalization and earlier sexual maturity.

It was reported "On average, the domestic foxes respond to sounds two days earlier and open their eyes one day earlier than their non-domesticated cousins. More striking is that their socialization period has greatly increased. Instead of developing a fear response at 6 weeks of age, the domesticated foxes don't show it until 9 weeks of age or later. The whimpering and tail wagging is a holdover from puppyhood, as are the foreshortened face and muzzle. Even the new coat colours can be explained by the altered timing of development. One researcher found that the migration of certain melanocytes (which determine colour) was delayed, resulting in a black and white 'star' pattern."


Archaeology has placed the earliest known domestication approximately 30,000 BC,[1][2] and with certainty at 7,000 BC.[3] Other evidence suggests that dogs were first domesticated in southern East Asia.[24]

Due to the difficulty in assessing the structural differences in bones, the identification of a domestic dog based on cultural evidence is of special value. Perhaps the earliest clear evidence for this domestication is the first dog found buried together with human from 12,000 years ago in Israel and a burial site in Germany called Bonn-Oberkassel with joint human and dog interments dating to 14,000 years ago.[25][26][27][28][29]

In 2008, re-examination of material excavated from Goyet Cave in Belgium in the late 19th century resulted in the identification of a 31,700 year old dog, a large and powerful animal who ate reindeer, musk oxen and horses. This dog was part of the Aurignacian culture that had produced the art in Chauvet Cave.[30][31]

33,000-year-old skull of a domesticated canid from Siberia

In 2010, the remains of a 33,000 year old dog were found in the Altai Mountains of southern Siberia.[32][33] DNA analysis published in 2013 affirmed that it was more closely related to modern dogs than to wolves.[34] In 2011, the skeleton of a 26,000 to 27,000 year old dog was found in the Czech Republic. It had been interred with a mammoth bone in its mouth—perhaps to assist its journey in the afterlife.[30][35]

Domestication of the wolf over time has produced a number of physical or morphological changes. These include: a reduction in overall size; changes in coat colouration and markings; a shorter jaw initially with crowding of the teeth and, later, with the shrinking in size of the teeth; a reduction in brain size and thus in cranial capacity (particularly those areas relating to alertness and sensory processing, necessary in the wild); and the development of a pronounced “stop”, or vertical drop in front of the forehead (brachycephaly). Certain wolf-like behaviors, such as the regurgitation of partially digested food for the young, have also disappeared.[36]

DNA evidence

The development of molecular biology allows us to infer evolutionary relationships for species and to represent them in a phylogenetic tree, however these are not without their limitations.

There are three schools of thought on the origin of the dog.

The Dog is a descendant of other canids that exist today

Within the Canidae, three distinct phylogenetic groupings are apparent. (1) the fox-like canids, which include species closely related to the red fox (genus Vulpes), as well as the arctic and fennec fox (genus Alopex and Fennecus, respectively); (2) the wolf-like canids including dog, wolf, coyote, Ethiopian wolf or Simien jackal, and three other species of jackals (genus Canis), as well as the African hunting dog (genus Lycaon) and the dhole (genus Cuon); and (3) the South American canids including fox-sized canids such as the pampus fox, crab-eating fox, and small-eared dog (genus Pseudolopex, Lycolopex, Atelocynus) and the maned wolf (genus Chrysocyon) and bushdog (genus Speothos). Additionally, there are several canids that have no close living relatives and define distinct evolutionary lineages such as the gray fox (genus Urocyon), the bat-eared fox (genus Otocyon), and the raccoon dog (genus Nyctereutes). These phylogenetic relationships imply that the dog has several close relatives within its genus, in fact, all members of Canis can produce fertile hybrids and several species may have genomes that reflect hybridization in the wild.[37] Therefore, it was once thought possible that the dog could have been a descendant of a number of canids that exist today - similar to the Coyote-Wolf-Dog hybrid found among some Eastern Coyotes[38] - until DNA analysis indicated otherwise.

The Dog is a descendant of a wolf that exists today

In 1995, DNA analysis of extant (i.e. living today) Canidea species indicated that the wolf and the domestic dog were so genetically similar that the wolf may have been the ancestor of the dog.[25] By 1999, further genetic evidence supported this view and indicated that the domestic dog may have emerged from multiple wolf populations.[39][40]

Based on these genetic indications, the domestic dog was reclassified as of 2005[41] as Canis lupus familiaris, a subspecies of the Gray Wolf Canis lupus. However, canis familiaris is also accepted.

The Dog is a descendant of a wolf that no longer exists today

During the Late Pleistocene megafaunal extinctions, a number of wolves disappeared that were genetically unique and morphologically distinct. In 2007, a study found that none of the 16 mtDNA haplotypes recovered from a sample of 20 Pleistocene eastern-Beringian wolves was shared with any modern wolf.[42]

In early 2013, a study isolated ancient DNA from the 33,000-year old skull of a dog-like canid that was excavated from Razboinichya Cave in the Altai Mountains of southern Siberia in 1975. The study then analysed 413 nucleotides of the mitochondrial control region. The analyses revealed that the unique haplotype of the Altai dog was "more closely related to modern dogs and prehistoric New World canids than it is to contemporary wolves".[4]

In late 2013, a study analysed the complete mitochondrial genomes from present-day dogs and wolves, as well as 18 fossil canids dating from 1,000 to 36,000 years ago from the Old and New Worlds. The data suggest that an ancient, now extinct, central European population of wolves was directly ancestral to domestic dogs. Furthermore, several ancient dogs may represent failed domestication events, such as the Goyet specimen of Belgium and the Altai Mountain specimen of Russia.[43][44][45]

In 2014, a study of the high-quality genome sequences of three gray wolves - one from each of three putative centers of dog domestication - two ancient dog lineages (Basenji and Dingo) and a golden jackal as an outgroup, indicated that "none of the wolf lineages from the hypothesized domestication centers is supported as the source lineage for dogs, and that dogs and wolves diverged 11,000–16,000 years ago in a process involving extensive admixture and that was followed by a bottleneck in wolves....and the sampled wolves instead form a sister monophyletic clade...This result, in combination with dog-wolf admixture during the process of domestication, suggests that a re-evaluation of past hypotheses regarding dog origins is necessary."[46][47][48]

Other Background information

mtDNA evidence suggests that African dogs, such as the Basenji, have a high level of genetic diversity.

Based on this DNA evidence, most of the domesticated dogs were found to be members of one of four groups. The largest and most diverse group contains sequences found in the most ancient dog breeds, including the dingo of Australia, the New Guinea Singing Dog, and many modern breeds, like the collie and retriever. Other groups such as the German Shepherd Dog showed a closer relation to wolf sequences than to those of the main dog group, suggesting that such breeds had been produced by crossing dogs with wild wolves. It is also possible that this is evidence that dogs may have been domesticated from wolves on different occasions and at different places. Vilà is still uncertain whether domestication happened once–after which domesticated dogs bred with wolves from time to time–or whether it happened more than once.

A 2002 study by Peter Savolainen et al. identified mitochondrial DNA evidence suggesting a common origin from a single southern East Asian gene pool for all dog populations.[24] In 2010, a study by Bridgett vonHoldt et al., using a larger data set of nuclear markers, pointed to the Middle East as the source of most of the genetic diversity in the domestic dog and a more likely origin of domestication events.[49] Z-L Ding et al. (2011) presented new Y-chromosome data from 151 dogs sampled worldwide, again pointing to a single domestication region south of the Yangtze river. The convergence of mtDNA and Y-chromosome variability in this region, the authors propose, indicate that a large number of wolves were domesticated, probably several hundred, as part of a sustained cultural practice.[50]

The most puzzling fact of the DNA evidence is that the variability in molecular distance between dogs and wolves seems greater than the 10,000–20,000 years assigned to domestication. Yet the process and economics of domestication by humans only emerged later in this period in any case. Based upon the molecular clock studies conducted, it would seem that dogs separated from the wolf lineage approximately 100,000 years ago. While evidence for fossil dogs lessens considerably beyond 14,000 years ago and ending 33,000 years ago, there are fossils of wolf bones in association with early humans from well beyond 100,000 years ago.[51]

Tamed wolves might have taken up with hunter-gatherers without changing in ways that the fossil record could clearly capture. The influx of new genes from those crossings could very well explain the extraordinarily high number of dog breeds that exist today, the researchers suggest. Dogs have much greater genetic variability than other domesticated animals, such as cats, asserts Vilà.

Once agriculture took hold, dogs would have been selected for different tasks, their wolf-like natures becoming a handicap as they became herders and guards. Molecular biologist Elaine Ostrander is of the view that "When we became an agricultural society, what we needed dogs for changed enormously, and a further and irrevocable division occurred at that point."[52] This may be the point that stands out in the fossil record, when dogs and wolves began to develop noticeably different morphologies.

A 2009 study of African dogs found a high level of mtDNA diversity. The authors suggest that a new view of the domestication of the dog may be needed.[53] A study by the Kunming Institute of Zoology found that the domestic dog is descended from wolves tamed less than 16,300 years ago south of the Yangtse river in China.[54] An older report said that all dog mitochondrial DNA came from three wild Asian female wolves.[55]


The Bulldog is well known for its short muzzle and saggy skin on its face
White Standard Poodle.
The fad for intentionally cross-bred dogs, such as this poodle hybrid, has resulted in the label "designer dogs"
A hunter with a large pack of beagles, a breed of hunting dogs, 1885.

As humans migrated around the planet, a variety of dog forms migrated with them. The agricultural revolution and subsequent urban revolution led to an increase in the dog population and a demand for specialization. These circumstances would provide the opportunity for selective breeding to create specialized types of working dogs and pets. These specializations originated simply from human selection for tameness alone.

Neoteny in the rapid evolution of diverse dog breeds

This rapid evolution of dogs from wolves is an example of neoteny or paedomorphism. As with many species, young wolves are more social and less dominant than adults; therefore, the selection for these characteristics, whether deliberate or inadvertent, is more likely to result in a simple retention of juvenile characteristics into adulthood than to generate a complex of independent new changes in behavior. (This is true of many domesticated animals.)

This paedomorphic selection naturally results in a retention of juvenile physical characteristics as well. Compared to wolves, many adult dog breeds retain such juvenile characteristics as soft fuzzy fur, round torsos, large heads and eyes, ears that hang down rather than stand erect, etc.; characteristics which are shared by most juvenile mammals, and therefore generally elicit some degree of protective and nurturing behavior cross-species from most adult mammals, including humans, who term such characteristics "cute" or "appealing". It has been seen that these traits can even prompt an adult female wolf to act more defensively of dog puppies than of wolf puppies.[56]The example of canine neoteny goes even further, in that the various breeds are differently neotenized according to the type of behavior that was selected.[57]

  • Herding dogs exhibit the controlled characteristics of hunting dogs. Members of this group, such as Border Collies, Belgian Malinois and German Shepherds, use tactics of hunter and prey to intimidate and keep control of herds and flocks. Their natural instinct to bring down an animal under their charge is muted by training. Other members of the group, including Welsh Corgis, Canaan dogs, and Cattle dogs, herd with a more aggressive demeanor (such as biting and nipping at the heels of the animals) and make use of body design to elude the defences of their charges.
  • Gun dog breeds used in hunting—that is, pointers, setters, spaniels, and retrievers—have an intermediate degree of paedomorphism; they are at the point where they share in the pack's hunting behavior, but are still in a junior role, not participating in the actual attack. They identify potential prey and freeze into immobility, for instance, but then refrain from stalking the prey as an adult predator would do; this results in the "pointing" behavior for which such dogs are bred. Similarly, they seize dead or wounded prey and bring it back to the "pack", even though they did not attack it themselves, that is, "retrieving" behavior. Their physical characteristics are closer to that of the mature wild canine than the sheepdog breeds, but they typically do not have erect ears, etc.
  • Scenthounds maintain an intermediate body type and behavior pattern that causes them to actually pursue prey by tracking their scent, but tend to refrain from actual individual attacks in favor of vocally summoning the pack leaders (in this case, humans) to do the job. They often have a characteristic vocalization called a bay. Some examples are the Beagle, Bloodhound, Basset Hound, Coonhound, Dachshund, Fox Hound, Otter Hound, and Harrier.
  • Mastiff-types are large dogs, both tall and massive with barrel-like chests, large bones, and thick skulls. They have traditionally been bred for war, protection, and guardian work.
  • Bulldog-types are medium sized dogs bred for combat against both wild and domesticated animals. These dogs have a massive, square skull and large bones with an extremely muscular build and broad shoulders.
  • Terriers similarly have adult aggressive behavior, famously coupled with a lack of juvenile submission, and display correspondingly adult physical features such as erect ears, although many breeds have also been selected for size and sometimes dwarfed legs to enable them to pursue prey in their burrows.

The least paedomorphic behavior pattern may be that of the basenji, bred in Africa to hunt alongside humans almost on a peer basis. This breed is often described as highly independent, neither needing nor appreciating a great deal of human attention or nurturing, often described as "catlike" in its behavior. It too has the body plan of an adult canine predator.

Of course, dogs in general possess a significant ability to modify their behavior according to experience, including adapting to the behavior of humans. This allows them to be trained to behave in a way that is not specifically the most natural to their breed; nevertheless, the accumulated experience of thousands of years shows that some combinations of nature and nurture are quite daunting, for instance, training whippets to guard flocks of sheep.


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Further reading

  • Klütsch CFC, Seppälä EH, Lohi H, Fall T, Hedhammar Å, Uhlén M, Savolainen P (2010) Regional occurrence, high frequency but low diversity of mitochondrial DNA haplogroup d1 suggests a recent dog-wolf hybridization in Scandinavia. Animal Genetics, online early.
  • Morey, Darcy. Dogs: Domestication and the Development of a Social Bond (Cambridge University Press; 2010) 384 pages; uses zooarchaeology to explores ties between humans and canines over the past 15,000 years with a focus on the New World and Arctic regions.

External links

  • Brisbin Jr., I.L.; T.S. Risch (15 April 1997). "Primitive dogs, their ecology and behavior: Unique opportunities to study the early development of the human-canine bond.". Journal of the American Veterinary Medical Association 210 (8): 1122–6. PMID 9108912. 
  • Trut, L.N. (1999). "Early Canid Domestication: The Farm-Fox Experiment". American Scientist 87 (2): 160–169. doi:10.1511/1999.2.160. 
  • Mestel, Rosie (October 1, 1994). "Living World: Ascent of the Dog".