a Single Origin to the Dog in Southern China 5.400 to 16.300 Years Ago
Summary by Dr.
© de Caprona
Arctic Wolf (Canada)
lupus arctos) © R. Schmode ~
lupus lupus) © G.Volker
~ Arabian Wolf (Canis
lupus arabs) critically endangered © S. Rerucha
photographs copyrighted to their photographer. Please do not use for
purpose without asking.
In their article
"mtDNA Data indicates a Single Origin for Dogs South of the Yangtze
River, less than 16,300 Years Ago, from Numerous Wolves", Pang et
(2009) analyse the mitochondrial DNA (mtDNA) of a large sample of dogs
to determine when, where and how the wolf was domesticated to become
The study of samples
ancient dogs have shown that ancient and modern dogs share the same
and that American dogs descend from the Old World dogs, hence the focus
of this study on the Old World dogs.
An indepth survey
patterns of geographical diversity for dog mtDNA was carried out by
582bp of the Control Region for 1.543 domestic dogs across the Old
Because it had been earlier indicated that the origin of dogs was
in East Asia (Savolainen et al 2002), this area was densely
This study includes
from Europe (North and South): Britain, Scandinavia, Miscellaneous;
Asia:Israel, Iran, Turkey, Miscellanous; Africa: North Africa, Central
Africa, South Africa, Miscellaneous;South East Asia: Thailand, Vietnam,
Cambodia; North China (Heilongjian, Liaoning, Hebei, Shanxi); Central
(Shaanxi, Shichuan); South China (Guangdong, Guanhxi, Hunan, Guizhou,
Yunnan; Tibet (Quinghai and Nepal) and Hainansanya. Samples were taken
on dogs in rural areas as they were assumed to have little influx of
dogs or on dogs which belonged to breeds with known historical origins.
Most of the "non-breed" dogs had specialized morphology and were
for specific purposes (guard, pet, herding, meat), they were not
The wolf samples
for the 582bp region were from Afghanistan, Canada, China, Mongolia ,
Russia, Saudi Arabia, and Yugoslavia.
Based on this
survey, a subsample
of 169 dogs representative across mtDNA diversity were selected for an
analysis of almost the entire mtDNA genome.
5 of the 6 main phylogenic groups or clades: A,B,C,D,F and E. The study
focuses on the first 3, as the clades D, E,and F are rare,
restricted and could be derived from post-domestication wolf-dog
The 2 major dog
A and B, contained wolf haplotypes.
1) Clade A had 3
from wolves of North China and Mongolia, which differed by 1 or 2
from the closest dog haplotypes;
2) In Clade B, 4
were found- 2 from North Chinese and Romanian wolves were identical to
dog haplotypes, and 2 from Afghani and Yugoslavian wolves differed by 2
substitutions from the closest dog haplotypes.
The study of wolves
remains incomplete because of the extinction of wolves in large parts
Europe and southern Asia and for this reason cannot give any tangible
as to the diversity of wolf haplotypes at the time of domestication.
on comparing the distribution of genetic diversity among the dogs
across the world to determine the place and time of the domestication
Tamed Mongolian wolf
(Canis lupus chanco)
Eagle Festival, West Mongolia
the Old World dogs share a homogenous gene pool:
A, B, and C were found in every population,
from 97.4%-100% of all dogs.
proportions of individuals with these haplotypes was similar across the
of the haplotypes
(9 in clade A, 2 in clade B, 3 in clade C) were termed Universal
The UTs were very frequent in the western part of the Old World (West
Ural and Himalayas): 81.5% of dogs in Europe and Southwest Asia had a
which is the same as a dog in Asia.
The UTs were much fewer in East Asia - 54.2% of the dogs there have
and in the extreme Southeastern part of China (South of the Yangtze
40.8% of the dogs have UTs with a range from 18.4% to 54.7% among the
in the West, all other haplotypes apart from the UTs differ by a single
mutation from a UT. 98.7% of European dogs and 94.6% of Southwest Asian
dogs either had a UT or a UT derived haplotype (UTd). This means that
all the dogs in the West had a haplotype traced back to a haplotype
found in East Asia
In contrast, only 69.2% of East Asian dogs and 53.0% of dogs South of
Yangzte river had UTds.
The genetic diversity follows a gradient, from maximum levels at one
of the continent South of the Yangtze River, decreasing through East
and Eurasia to reach the lowest levels in Europe at the other end of
Clade A in particular, Western populations lacking several mostly
identical parts of Clade A, the populations having an almost complete
across clade A being those South of the Yantze River.
Western dogs have a haplotype pool which consists almost exclusively of
14 UTs and surrounding haplotypes. East Asian population particularly
Southern China have a large proportion of dogs with unique haplotypes a
large distance away from the UTs, therefore also from the haplotypes
in the West.
complete mtDNA genome analysis shows that clades A, B and C consist of
several subgroups. Clade A has 6 major subclades, Clade B and C have 2
subclades each, for a total of 10 subclades or haplogroups.
The complete set
subclades is found only in Southern Eastern China, 7 are found in
China, 5 in Northern China/India/SWAsia, and only 4 in Europe. 5 of the
6 Clade A subclades were found only in East Asia. In Southern China, no
province had all 10 subclades, but in Yunnan, Southeast Asia and
9,9, and 8 of these subclades were found. The smallest region
all 10 subclades is in the Southwest area
South of the Yangtze River.
substitution rate for the mtDNA data was calculated to estimate the age
of the phylognetic groups. The fossil record is incomplete and the
time for dog/wolf and coyote cannot be calibrated exactly. The data
indicate that clades A,B and C spread simultaneously across the Old
5.400 to 16.300 years ago.
The minimum number of lineages was estimated for the whole mtDNA genome
data which existed at the time of the spread of the dogs across the Old
World. Haplotypes separated by more than 12 substitutions from each
should have originated from different founders at the time of the
counting such haplotypes, the authors find a minimum of 51 different
wolf lineages behind today's dogs' haplotypes. This estimate is
and the authors assume that several hundred female wolves were
They base their assumption on the fact that
differing by less than 12 substitutions could be from different female
may be present that are not detected in this data set
may have become extinct since the domestication of the wolf
wolves may have had the same haplotype
The exact area South of the Yangtze where the domestication of the dog
may have taken place is not clear. The full representation of all 10
is found in a region which includes Yunnan, and South East Asia.
has more haplotypes than Yunnan and only one subclade less.
simplest explanation for the geographical distribution of the 10
of clades A, B and C is that they had a single origin within the area
to the South of the Yangtze river and South Eastern Asia. From there a
subset of the original gene pool spread to the rest of the world.
of these 3 clades across populations of the old World indicate that
populations originate from a founder population with a similar
of the 3 clades. Had the domestcation of the dogs happened
in various areas, one would expect a large proportion of regionally
haplotypes and varying proportions of the clades would be expected.
showed that multiple origins in time would have required extreme
rates and thorough mixing of the populations to end up with the
proportions of the clades, an unlikely event. The only scenario which
moderate migration is that of a single origin in time and space for the
is not possible to date the origins of dogs from these data, but the
of the spread from the center of origin can be estimated. If the dog
spread shortly after the domestication of wolves, the genetic data
that dogs originated approximately 5.400 to 16.300 years ago.
in the mtDNA genome corresponds to 3.200-9.600 years, the sharing of
and UTs across Eurasia is not caused by recent mixing of populations
results from ancient events.
results show for the first time reasonable estimates for the time of
of dogs and the number of female founder wolves. They dismiss the
by Vila et al 1997 that mtDNA indicate an origin of dogs much earlier
10.000 to 15.000 years ago.
The European dogs lack 6 of the 10 subclades. They have a special
compared to other dog populations in that they have specialized in a
large number of dog breeds. For this reason one could think that the
genetic diversitry of European dog breeds results from severe
during the development of the many different breeds. However the gene
is almost identical to that of the dogs from South West Asia. Therefore
the 6 subclades must have been missing already before the development
the many different breeds in Europe. In spite of the largest
variation in all of their many breeds, the European dogs have the
genetic variation, and this fact places them at the periphery of the
dog populations, and does not reflect modern breeding history.
The analysis of the whole mtDNA genome, as opposed to the mtDNA bp
brings the necessary finer resolution needed to time the dog
more appropriately in good agreement with archaelogical finds.
It is possible that the domestication of wolves was a widespread
which took place in various regions South of the Yangtze River. The
Chinese dogs' genetic diversity is very low and comparable to the
dogs with only 5 subclades and 89.8% UTds. The wolf was present in
Eastern China until the 1950s although it is now extinct there.
Olsen and Olsen (1977) found that a morphological feature typical of
dog's jaw is found also in Chinese wolves ( the turned-back apex of the
coronoid process of the ascending ramus). Added to the relatively small
size of the Chinese wolf, these 2 authors had concluded that the
wolf was a likely ancestor of the dog.
This study focuses only on the mtDNA of today's dogs and therefore only
on the domestication of the female wolves which were ancestral to
If there was any
or domestication of wolves or other canids which are not ancestors of
dogs, this study does not reveal it.
give for the
first time some indication as to which human culture may have tamed the
wolves. The time and place of the domestication of the wolves coincides
with the transition from hunting to farming and the emergence of rice
study shows that a large number of wolves were domesticated suggesting
that the domestication was not a chance/isolated event but probably a
custom - it also shows that the domestication of wolves happened only
that the wolf was first tamed as a source of food, as dogs have been
on a large scale in Southern East Asia until today. They suggest that
use of dogs for herding, hunting or companionship was perhaps the
of selection by European non-dog eating peoples.
The authors state
thorough studies in South East Asia of archaelogical canid remains, and
of genetics of modern and ancient canid samples should help to
the validity of this hypothesis. The authors also stress that a similar
study on the male lineages (Y chromosome) would complement this study
the mitochondrial DNA of female wolf and dog lineages (mtDNA being only
inherited through female lines)
Peter Savolainen and Arman Ardalan for their positive comments,
Bologov for the Russian wolves picture, Simon Rerucha for the Arabian
picture, Alexander Choi Jackson for his picture of the
Wolf and boy, and Philippe Touret for providing
invaluable pictures of Chinese dogs and the information pertaining to
Olsen JW (1977): "The
Chinese Wolf, Ancestor of New World Dogs", Nature 197: 533-535
Kluetsch C, Zou
XJ, Zhang AB, Luo LY, Angleby H, Ardalan A, Ekström C,
A, Lundeberg J, Matsumura S, Leitner T, Zhang YP, Savolainen P.(2009):"MtDNA
Single Origin for Dogs South of Yangtze River, less than
16,300 Years Ago, from Numerous Wolves." Mol Biol Evol. 2009
Luo J, Lundeberg J, Leitner T (2002): "Genetic evidence for an
Asian Origin of Domestic Dogs." Science 298:1610–1613.
dogs, photographed in the Guangxi and Guizhou areas, South of the
River in August of 2009
(none of these dogs
strays, they all belonged to a specific house and someone who took care
of them; the information in the photos' captions is given by Philippe
Quan" in the small horned Miao*
of Boji. The breed, which is named after the village of Xiasi,
now accepted in most Chinese dog shows.
Matang village of the Gejia people (Guizhou). All the dogs of this area
were taller, more powerful and long-legged and of that off-white
In a collection of
of pictures of this region's dogs, the first time these dogs are
small town of Yangxi (Guangxi)
Front :a "Pig-Dog"
minority Dong. Its name relates to the fact that once it has been
slaughtered and skinned,
this dog looks like
its body being covered with a skin of fat.
Back: a typical
dog of the Guangxi region. The curled tail is not the tail carriage
by the local hunters
who prefer a
carried high, which enables them to see the dog in high grass.
(Guizhou), village inhabited by black Miaos*.
The local dogs
an intermediate type between the dog types of Guangxi and Guizhou
(Guizhou), only village during that trip where brindled dogs were
The puppy resembles
Chow-Chow, his mother has the characteristics of an official Xiasi
except her color.
The white color for
breed is imposed by the standard but other coat colors can be found (
fawn, slate blue, brindle)
in small horned Miao* village of
(Guizhou). Type close to the breeds hunting boar in neighboring Guangxi.
Touret 2009 ~
© Jean-Maurice Touret 2009
(Guizhou). Dog of original Chow-Chow type, roots of this
breed to be found towards Guangdong
Miao village of Gaoyao (Guizhou).
Dog close to the
type, slightly wire haired,
for the region where solid colors are the absolute majority.
black Miao* village of Biasha
© Philippe Touret 2009
Miaos are a group of linguistically related people in Southwest China.
They are referred to in Chinese as "small horned" or "long-horned"
depending on the length of the horns of the headpiece they wear, or
white, red, striped, small flowery, big flowery, green, blue Miaos
on the most characteristic color of the women's clothes.