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This gampr in Armenia may reflect the consistent breeding in the beginnings of the domestication age, where crossing with wolves was regularly practiced. Photo by Hamlik P.

DOMESTICATION

 

Pre-History

around the world

The location of the first domestication of the dog seems to change depending on the method of investigation.  What has become apparent however, is that the original ancestor of all dogs was not exactly a dog, nor was it the same as any wolf living now; it was what may be called a proto-canid and lived until somewhere between 10,000-35,000 years ago.

The proto-canid has some very interesting descendants in groups of non-domesticated dogs in various locations around the world. These non-domesticated dogs have been referred to as pariahs, when in actuality they are dogs who were never domesticated, but in some cases acclimated to living among humans.

Some of these breeds of non-domesticated dogs are:

 

These breeds can live entirely without human assistance. Most ovulate once per year, are able to live alone or in packs, and all can be domesticated.  Typical features include upright ears, almond shaped eyes, short coat that is most commonly golden/tan in color, tapering muzzle, and many have a curled tail.

By mtDNA analysis, the dog species which has changed the least from the original proto-canid is the New Guinea Singing dog. These dogs are nearly extinct in the wild. They have a unique style of vocalizations and may be the most difficult to adjust to human cohabitation.

Many scientific studies of dog domestication seek to prove a single point of origin. The lens though which most professors see the process is that the domestication incident of a dog happened in one location; and from that location then spread to other areas; and that all dogs are descended from that one incident.

Various locations have been proposed and given scientific credence, including Southeast Asia, Mesopotamia, Siberia and Europe.

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Indian Native Dog, the InDog, working as a Livestock Guardian in India

However, when we examine the localized populations of non-domesticated dog species, such as those listed above, we see multiple spontaneous incidences of the dog-human partnership continuing even in modern times. It has been documented in Israel, India, Australia and various places in Malaysia, plus southeastern USA, in a population of wild dogs that has been disconnected from the others for over 10,000 years.

So to propose that dog domestication stemmed from a single incident rather than multiple incidents around the world is failing to look at the likelihood that the dog-human partnership is so natural and useful, it is an eventual foregone conclusion.

Furthermore, among the wild purebred dog species who have occasionally adapted to cohabitation with humans, various skills seem to be inherent; one such skill is an occasional natural inclination to work as a ‘livestock guardian,’ which requires a very complex set of abilities and instincts.  Individual dogs in India and in Israel can be found guarding livestock. The act is well documented. The dogs can also perform other functions.

The InDog is nearly identical to the Canaan dog of Israel. They also very much resemble the Australian Dingo. The dogs, and others that Rajashree Khalap has documented, are filling the function of Livestock Guardian dog.

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In Israel, the Canaan dogs also have been known to work with livestock

The comparison of these wild dogs, in a common "buff" color type, from far reaches of the world. In the latter half of the gallery above, the  last six photos are near-wild dogs who have become habituated to living with humans, but have not been changed much by domestication.

The strong similarities among the dogs above are clear. These dogs are from several continents, and have not interbred in centuries, if not millennia. All show a strong phenotype similarity, and because they are genetically more similar to the ancient proto-canid ancestor of all dogs, we can assume that they do represent some sort of basic dog, not affected by domestication so much as merely the necessities of survival.

Analyzing the effects of the domestication process, most clearly presented in the Trut fox study (see below), we can see that the natural results of domestication (selection for friendliness toward humans) are what can also be seen in nearly all species of domesticated animals.

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Canaan Dog puppies - white points advancing to cover base color

 

Selection study

phenotype changes

Fifty years ago, Dr. Dmitry Belyaev and his team at the Institute of Cytology and Genetics in Siberia pioneered a strict study of domestication in foxes. Beginning with 130 foxes, the team permitted only specific, timed, consistent interactions with the captive foxes, and allowed only a small percentage of them to be bred. Selection was based only on friendliness, and disregarded any other means of distinguishing those which were allowed to reproduce. A remarkable progression of developmental and physiological changes were recorded by the team, even after the death of Dr. Balyeav.

Analyzing the effects of the domestication process, most clearly presented in the Trut fox study, we can see that the natural results of domestication (selection for friendliness toward humans) are what can also be seen in nearly all species of domesticated animals.  When observing characteristics of domesticated animals, it becomes apparent that the process of domestication is linked to certain physiological traits (Trut, Lyudmila N., March-April, 1999) which progress in a surprisingly consistent manner for all species.

With selection being based only on friendliness, the speed of the domestication process was dramatically increased, accomplishing inadvertently various developments in fifty years which had never been done, even with intention of doing so, in the history of the farm-fox business (Trut, 1999). Apparently the selection factor of friendliness is polygenic, bringing with it certain traits which can be observed in many other breeds of livestock. Three main physiological traits expressed themselves early and consistently. White pigmentation on first the head then the feet, flopping ears, and curling of the tail were all evident in the early stages of domestication.

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Dr. Lyudmila Trut with a domesticated silver fox in Siberia Photo courtesy of Dan Child/©BBC

Developmental changes were remarkable as well. Dramatically, the basal levels of corticosteroids in the blood plasma of the domesticated foxes had dropped to slightly more than half the level in a control group by just 12 generations, and within another six generations had halved again (Trut, 1999) Lower corticosteroid levels indicate a lower level of fear, and also less energy usage in order to produce the fear reaction. With this comes a higher level of serotonin, which in turn affects neonatal development. These changes affect the timing of certain growth markers, such as “earlier eye opening and response to noises and the delayed onset of the fear response to unknown stimuli.” (Trut, 1999).  These in turn affect an animal’s ability to be aware of and to accept interaction with humans.  Other significant early changes in morphology were differences in skull proportions, more variety in fur color, and changes in size.

As domestication progresses, selection being based on a criteria arbitrarily selected by the desire of humans rather than functional need, many non-productive traits crop up. The Belyaev team began to see malformations of the jaw structure, short legs, bowed legs, extreme desire for closeness with humans, and less difference in size between the males and females (Trut, 1999). If the foxes had been living in a situation where their survival, and therefore ability to reproduce, depended on being physically functional, it may be safe to assume that some of these less desirable characteristics would not have become prevalent. However, since the study illustrates the connection between friendliness and these morphological differences, one could also surmise that the domestication of the foxes in a more normal environment would have also been halted or dramatically slowed at the point of non-functionality.

 

Wolfy genetics?

proto canid

Until recently, the common assumption was that dogs are descended from domesticated wolves, most likely the common gray wolf. There are many references to show a genetic relationship, but, the leap from wolf to dog, specifically, was difficult to pinpoint. Until the discovery of the proto-canid, that is.

Viewed from the perspective that all domesticated dogs are descended from the proto-canid species that may have been ubiquitous, or became so throughout the millenia, we can easily see how intermittent mixing with wolves throughout history has affected populations of dogs in various places.

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(Left) Steppe Wolf, (Right) Iranian wolf (canis pallipes)

(Below) Caspian Sea wolves

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Recent mtDNA analysis of dogs selected within the modern boundaries of Georgia found that the native livestock guardians have ~23% local wolf genes, and the local wolves sampled had ~12% dog genes, indicating a significant level of natural crossbreeding between the Caspian wolf and local guardian dogs. Disturbingly, this article and others like it completely fail to mention the existence of Armenia except when absolutely unavoidable, in maps, as if Armenia does not exist. 

According to a very thorough analysis published in PLOS Genetics,

the genetic exchange/mixture with various populations of wolves has happened intermittently and in various locations throughout the last ~15,000 years. Prior to this time, a genetic bottleneck occurred, reducing the diversity of Canus to a fraction of the previous populations.  The bottleneck, and subsequent various descendants, are illustrated in the graph below.

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The study analyzed various wolves, (although not the Caucasus wolf, so specific data for that species is still unknown), a boxer dog, basenji, dingo and Golden Jackal.

The graphs on right side of the image show divergence of the populations after the genetic bottleneck, with the horizontal bands representing admixture during various spans of time. As you can see, the basenji and the Israeli's wolf intermixed a few thousand years ago, and the dingo and Chinese wolf intermixed prior to the dingo becoming isolated on the Australian continent.  To read the charts correctly, please visit the article at PLOS Genetics.   To summarize however,  localized Proto-canid descendants) + (various localized species of wolves) + (genetic changes due to developing relationship with humans) = aboriginal domestic dog.

Because of the ubiquitous nature of the wild dog varieties, and the presence of several species of wolves from distant locations, within the genetic history of our domestic dogs, it is not logical to assume that domestication began as one single incident.  As is said in the Reterieverman Blog, "Now, it doesn’t mean that phenotypically distinct domestic dogs first appeared in the Middle East. It just means that this population contributed to the majority of dogs we have today. The small dog gene that is found in purebred dogs is also found in some Middle Eastern wolves, likely coming from Canis lupus arabs. The smallest members of that subspecies weigh only 25 pounds.”

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Images by Ani Hayaserian, found here.

 

Mesopotamia

cradle of civilization

Most farmers know, the best deterrent for predation of livestock is a Livestock Guardian Dog, or LGD. As Armenians know, it is nearly impossible to keep sheep safe without a pack of dogs for protection. For these domestication events to have taken hold and lasted, the livestock must have been protected in some way. Of course, there were none of our modern methods available at the time which leaves us with the only logical conclusion that the livestock were protected by dogs.

If we look at an overview of domestication of all animals, this gives a strong bias for some domestication event in Mesopotamia:

  1. Dogs were the first domesticated animal, location(s) not clear

  2. Sheep and goats were first domesticated in Mesopotamia

  3. Followed by pigs in Mesopotamia

  4. Then cattle in Mesopotamia, with a second and separate domestication event in Pakistan

  5. Horses were then domesticated in multiple areas from different strains of wild horse (giving credence to the concept of multiple domestication events for dogs also)

  6. Llamas were domesticated in South America ~4,500 years ago

The two artifacts below are from Mesopotamia dated between 500-2000BCE.  Both are wearing a collar which is a sign of domestication (not just cohabitation), and they show a distinct similarity to the closest descendants of the proto-canid, as pictured above.​

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(Left) Sumerian dog Pendant, (Right) Elamite Dog Amulet - Goddess Gula, healing

This type of figurine is numerous in Armenian archeological finds. These pendants are much more fierce and wolf-like than the dogs above; but they are not fully wolves, as can be seen by the small curl at the end of their tails.  These figures have a much larger and heavier cranium compared to the dog figures above, however, they still have erect ears.  These are likely be representative of a stage of development of the gampr.

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Wolf-dog figurines, found near Ayrum in Shirak, dated 600BCE, Gevork Nazaryan

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Voski, is strikingly similar to the wild descendants of the proto-canid, but much larger as would be needed for protection of sheep

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This photo, almost 100 years old, taken in Nakhijevan. The dogs are more typical of a proto-canid descendant that also had some local wolf in its ancestry, Then carefully selected for the correct traits over the centuries