The study of dog domestication involves several different areas of expertise including the

work of zooarchaeologists, paleontologists, and evolutionary geneticists (genomicists). The

domestic dog may be one of humanities greatest triumphs as they are the only large carnivore
ever domesticated and the only domestic species to have created a unique social niche in human
society, other than the house cat. Descended from grey wolves, the dog was the first animal
domesticated by human beings tens of thousands of years ago, long before humans began to
experiment with domesticating other species of animals and plants for agriculture (Fox, 1978).
The duration of this process has been in debate for some time, but recent Russian Belyaev Fox
Farm studies on domestication have shown that it can occur in just a few successive generations.
Both the date and location of the events surrounding the first domestication of dogs has also been
debated for decades, as well as the method by which this occurred. While some like Dr. C. Vila
and Dr. Robert Wayne have claimed that domestication may have occurred as far back as
135,000 years ago until recently the oldest accepted domesticated dogs dated to around 14,000
years ago (Vila et al, 1997). Domestic canines are well known in the archaeological record for
the last 14,000 years, but there are far fewer that have been discovered that date before that time
(Ovodov, 2011). However, due to new discoveries, the oldest known possible dog or dog-like
canines remains were discovered in Horizon 4 at the Upper Paleolithic site of Goyet in Belgium
during the 19th century. In 2008 the remains were AMS radiocarbon dated to 31,700 or 36,000
BP (Ovodov, 2011). This AMS date was arrived at by direct dating of the remains themselves,
since much of the contextual data had been lost in the century since the excavations. These
skeletal remains were morphologically different from wolves, but not consistent with all the
features of modern dogs and are currently referred to as an incipient dog (Ovodov, 2011). This,
however, is not the only evidence of dogs or dog-like canines to be dated older than the accepted
14,000 BP; skeletal remains have also been recovered in Razboinichya Cave (Altai Mountains),
Russia in 2011 which date to 33,000 BP and from Predmost in the Czech Republic (three
specimens) which could be as old as 27,000 years old (Germonpr et al, 2012). The footprints
made by a child and a dog, or domestic wolf, were found at Chauvet Cave, France which date to
26,000 BP, but this is still very controversial
(http://www.nbcnews.com/id/27240370/ns/technology_and_science-science/t/worlds-first-dog-
lived-years-ago-ate-big/#.U2RpuRUo7IU). These discoveries challenge the notion that
domestication occurred only by or after 14,000 BP, but the small number of these older remains
leaves open the question of when and where humans first domesticated the wolf and if this was a
centralized process beginning in a single region or multiple events that occurred over time and
space. There have been many archaeological discoveries and genetic studies concerning the
domestication of dogs, for this paper I will focus only on the most recent.
Archaeological analysis
Archaeologically the study of dog domestication has revolved around the study of the
morphological differences (osteometrics) seen in the skeletal remains of ancient dogs recovered
from human occupation sites and then compared to modern dogs and their wild ancestors, the
wolf. These morphological changes in the skeletal system include a reduction in overall size,
smaller skull size averaging 20% smaller, a shorter jaw initially with tooth crowding and, later,
the teeth themselves shrink in size, and the development of a vertical drop in front of the
forehead known as brachycephaly (http://www.pbs.org/wnet/nature/episodes/dogs-that-
changed-the-world/photo-essay-from-wolf-to-dog/1278/.) All these differences translated into
different average measurements which are no longer consistent with those of the grey wolf.
Measurements are taken from the recovered skeletal remains and compared to known values
associated with the range of variation for modern and ancient dogs and wolves. These
measurements allow researchers to know which category to place the remains in. This process is
complicated by the fact that very early specimens are still difficult to distinguish from wolves
(Larson et al, 2012). In living specimens there are other morphological changes including
changes in coat coloration and markings, as well as differences in the structure of the ears from
erect ears like those of wolves to the floppy ears seen on hounds and other domestic breeds.
Many of these phenotypic changes were seen firsthand in the recent Russian Belyaev Fox Farm
studies on domestication involving silver foxes which began in 1979 and continue to this day
(Braude and Gladman, 2013). As stated above the earliest AMS radiocarbon dated dog remains
are those from Goyet cave in Belgium and the remains discovered in the Altai region of Siberia.
Osteological and genetic analysis of the Altai remains, however, would seem to indicate that this
incipient dog, while strongly resembling the ancient Greenland dog, is not related to modern
domestic dogs and represents an early but ultimately unsuccessful attempt at domestication
(Ovodov, 2011). While this does not represent the true lineage of modern domestic canines it
does support the idea that domestication was attempted at numerous locations and at several
different times throughout prehistory (Ovodov, 2011).
However, assessing the morphological differences in the skeletal remains recovered from
archaeological sites is not the only method archaeologists have used in the study of the origins of
the domestic dog. Cultural evidence has also been used to identify some of the earliest definitive
domestic dogs. This evidence includes dog burials and more importantly joint human-dog burials
which are known in the archaeological records as far back as 14,000 BP at sites like Bonn-
Oberkassel in Germany in which a dog was interred in a human double grave
(http://archive.archaeology.org/online/interviews/morey.html). The second oldest site of joint
human-dog burials is Ein Mallaha in Israel, where an elderly human and a 4-5 month old puppy
were buried together (http://scienceblogs.com/thoughtfulanimal/2010/04/12/monday-pets-
biological-evidenc/). This human-dog burial has been dated to sometime between 10,000 and
12,000 BP (http://scienceblogs.com/thoughtfulanimal/2010/04/12/monday-pets-biological-
evidenc/). These burials are some of the most important forms of evidence because they give us
the remains necessary for osteometric and genetic analysis.
Burials are not the only cultural evidence of early domestic dogs though. Human made
artifacts may also give insights into the early relationship of humans and dogs. The side by side
footprints of a child and an early dog/domestic wolf that were found at Chauvet Cave, France
dated to 26,000 BP (http://www.nbcnews.com/id/27240370/ns/technology_and_science-
science/t/worlds-first-dog-lived-years-ago-ate-big/#.U2RpuRUo7IU). Other examples include
cave paintings and other rock art from the Neolithic, like that found at Tassili n Ajjer, Algeria,
and a range of artifacts depicting dogs made during and after this early period have been found
all over the world. These artifacts often show dogs after domestication and are less useful in
determining the date of initial domestication. However, these artifacts are useful in determining
the minimum date for the regional presence of domestic dogs and the possible antiquity of some
modern breeds.
Genetic Analysis
Over the years since the invention of various methods of DNA analysis there have been
many studies undertaken to determine when and where dog domestication first occurred. These
studies generally compare the mitochondrial genomes of archaeological specimens to modern
dog breeds and wild canids.
MtDNA was discovered in the 1960s by Margit M. K. Nass and Sylvan Nass. MtDNA is
found in the mitochondria which are the energy-producing organelles of a cell. These are present
in large quantities in animal cells and are far more likely to survive over time than nuclear DNA
which can become degraded. MtDNA is inherited from the maternal ancestor and has a
predictable rate of mutation and as a result it is the material used in the analysis to examine the
evolutionary change in human and non-human species. These mitochondrial DNA strands are
extracted, then sequenced, and finally compared to the MtDNA sequences of others of the same
species or to related species.
The results of the early genetic studies pointed to various areas around the Old World
including Europe, China, and the Middle East. Many of the early studies were flawed however
because they focused only on the genetic material of modern dog breeds. Other early studies, like
that of Dr. Vila, returned with dates as far back as 135,000-140,000 years ago (Vila et al, 1997).
However, no archaeological evidence discovered to date exists to support such early beginning to
canine domestication. More recent studies have suggested dates for domestication that are much
more consistent with the archaeological evidence, the finding of two of the most famous studies
are as follows.
In 2013, Dr. Olaf Thalmann conducted a studied of canine MtDNA to determine the
origin of modern domestic dogs. This study looked at ten ancient wolf-like animals and eight
dog-like animals recovered from archaeological sites that range in age from 1,000 years old to
33,000 years old (Thalmann, 2013). The MtDNA recovered from these animals was then
compared to modern mitochondrial genomes of 77 domestic dogs, 49 wolves, and four coyotes
(Thalmann, 2013). The conclusion of this study was that modern domestic dogs were not closely
related to the modern wolves used in the study but likely were descended from an extinct group
of wolves from Europe and that domestication may have begun there as early as sometime
between 18,800 and 32,100 years ago (Thalmann, 2013). This study is interesting because it
refutes the idea that domestication began in the Middle East as wolves were drawn to early
human settlements during the Neolithic Revolution.
Other studies like that undertaken by Freedman in 2014 on the genome sequencing of
domestic dogs indicates that modern domestic canines developed between 11,000 and 16,000
years ago, still predating the rise of agriculture and sedentism (Freedman, 2014). This study
compared the MtDNA of grey wolves from three distinct regions believed to be centers of
domestication, two Basal dog lineages, and a golden jackal (Freedman, 2014). They also
discovered a genetic bottleneck in wolves which occurred just after the separation of dogs,
indicating that dogs were developed from a much larger pool of wolves (Freedman, 2014). This
date range is well supported by the archaeological evidence which now suggests domestication
just prior to the development of agriculture, but this does not explain the much earlier dates from
some archaeological finds such as the Goyet dog. It does, however, give an explanation as to
why the incipient dogs genetics are so different from modern wolves.
In conclusion, recent studies utilizing modern methods have given us new insights into
the domestication of the dog. New discoveries such as the Altai and Goyet dogs may push the
date of domestication back thousands of years. However, this new information has not settled the
debate over where and when domestication occurred. It now appears that there may have been
multiple occurrences of domestication, some of which did not contribute to the genetics of
modern domestic dogs. These studies have also discovered a genetic bottleneck in wolves that
may account for the genetic differences in ancient dogs and modern wolves. This may aid
researchers in discovering the geographic origin of the domestic dog through further genetic
testing if MtDNA can be discovered from these lost lineages.
















Works Cited
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Fox, Michael
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London
Germonpra, Mietje, Martina Lznikov-Galetovb, Mikhail V. Sablinc,
2012 Palaeolithic dog skulls at the Gravettian Pedmost site, the Czech Republic
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Larson G, Karlsson EK, Perri A, Webster MT, Ho SY, et al.
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Ovodov ND, Crockford SJ, Kuzmin YV, Higham TF, Hodgins GW, et al.
2011. A 33,000-year-old incipient dog from the Altai Mountains of Siberia: evidence of
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Thalmann O. et al.
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Viegas, Jennifer
2008 World's first dog lived 31,700 years ago, ate big; Discovery could push back the
date for the earliest dog by 17,700 years
(http://www.nbcnews.com/id/27240370/ns/technology_and_science-science/t/worlds-first-dog-
lived-years-ago-ate-big/#.U2RpuRUo7IU)
Goldman, Jason G.
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(http://scienceblogs.com/thoughtfulanimal/2010/04/12/monday-pets-biological-evidenc/)
Archaeology
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(http://archive.archaeology.org/online/interviews/morey.html)


Nature; Episode Dogs That Changed the World
http://www.pbs.org/wnet/nature/episodes/dogs-that-changed-the-world/photo-essay-
from-wolf-to-dog/1278/