Dogs in Ancient China

Dogs in Ancient China

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Dogs are the oldest domesticated animal in China and were bred as guardians, for transporting goods, for herding, hunting, and as a food source. Archaeological evidence dates the domestication of the dog in China at approximately 15,000 years ago. Remains of dogs have been found in Neolithic graves and their bones in middens dating from the same period onwards.

In ancient China, the dog was always regarded along utilitarian lines as a worker, not as a companion. Even so, dogs were regarded highly not only for their practical uses but as liminal beings who bridged the span between the realms of the living and the dead, the mortal and the unseen world. Chinese folklore often features a dog with transformative qualities or one associated with the divine, and the dog serves as the 11th sign of the Chinese Zodiac.

The famous loyalty of the dog is emphasized not only in folktales and legend but also in art and protective iconography. Jade amulets were often carved in the form of dogs and dog statuary was placed outside of homes, businesses, and royal residences. The dog was recognized early on as a protective force and this image persists to the present day, even though now they are also valued as pets and members of the family.

Early History

The Neolithic site of Banpo Village in Shaanxi Province is one among many offering evidence of the dog's early domestication. The village was occupied between 4500-3750 BCE, and there is no doubt the residents kept many dogs during that time. Dog and pig bones have been found in abundance, and it is possible, even likely, that by this early date dogs were already associated with the spirit world. An image which resembles a dog has been found on ceramic shards along with that of the mythical pig-dragon (precursor to the now famous Chinese dragon and also representing good fortune and protection) suggesting the villagers recognized a spiritually significant aspect of the dog.

Primarily, however, the dog was a functional tool in the everyday life of the village. Scholar Judith M. Treistman writes:

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The dog was probably used for food and clothing but most significantly for transportation. In this latter capacity of draft animal, one which was important to later tribesmen as well, the dog was part of the technology of forest efficiency. (31-32)

Although the people of Banpo were primarily vegetarian, they hunted wild game and fished in nearby waters. Treistman notes that the animals hunted included wolf, polecat, mountain sheep, musk, deer, roe deer, wild boar, elk, weasel, and many others. Dogs were used to haul the dead animals back to the village for processing. Once they became too old to assist the hunters, they were most likely killed themselves for their coats and whatever other parts of their bodies could be used.

They were also killed to release their spirit for protection and were buried in front of homes, not only at Banpo but throughout later Chinese history. Ghost stories were the earliest form of literature in China and, as with virtually every ancient culture, the Chinese had a deep-seated fear of ghosts. The most common reason for a haunting was improper burial, and if a ghost appeared, it was usually for either this reason or that the deceased's tomb had been disturbed. There were many different kinds of ghosts, however, and some were simply ill-tempered or disturbed spirits whose interest was in possessing a person or making their lives miserable. Since dogs were recognized as natural protectors, it made sense that they would be especially effective against spirits as spirits themselves.

Dogs as Spiritual Defenders

In time, the practice of sacrificing dogs fell off, and straw dogs were placed before a home or a city's gates for the same purpose, and this observance eventually gave way to statues of lion-dogs, usually made of stone, and known as Foo-Dogs in the west (foo meaning Buddha). This concept originated when Buddhism came to China from India accompanied by the myth that Buddha would ride through the sky on the back of a lion. The Chinese had never seen an actual lion but they had seen the small dog known as the Pekingese. The Pekingese was bred and carefully groomed to look like the descriptions the Chinese had heard of the lion and the Foo-Dog statue, then, came to represent the protective power of the Buddha.

The Foo-Dog statue came to represent the protective power of the Buddha.

The stone beasts, usually growling and adorned with an ornate belled collar, served the same purpose as the corpse of an actual dog or a straw figure. The bell was thought to warn of the approach of a spirit while the collar was engraved with symbols for protection and, often, the head or face of a spiritual protective entity. They were always placed in pairs; the male has a paw on an ornate ball, representing power and domination over the human affairs of the world while the female has her paw on a wriggling cub, symbolizing control of the forces of nature.

The belled collar motif came from the practice of attaching bells to actual dogs' collars as it was believed that bells (or chimes) would ring in the presence of unseen spirits. The pairing of the Foo-Dogs is also significant in that the female represents the yin force and protects the people of the house while the male represents yang and protects the structure itself.

This exact practice is still observed today in China and elsewhere outside Chinese-owned establishments. One will frequently see lion-dog statues to either side of the door of a Chinese restaurant, and these are there to guard against evil spirits and dark forces. If one pauses to view the statues carefully, one will see that the cub beneath the female's paw appears to be suckling. This detail comes from the ancient belief that lionesses secreted milk through the pads of their paws. The lion-dogs do not resemble lions or dogs so much as mythical creatures from another world, and this, of course, is by design. Dogs were often associated with the in-between realm which separated life and death, which contributed to the high esteem they were held in even as they continued to be used as a source of food.

Dogs in Folklore

Dogs were considered so important, in fact, they were thought to deserve divine protection for their services. One folktale tells the story of a man named Coffin Head Li who was a cruel bully preying on helpless animals, especially dogs and cats. One day, Coffin Head is visited by two men in purple robes who tell him that he has been condemned by the gods of the afterlife for his abuse of animals. Coffin Head laughs their comments off as a joke but they then produce a document detailing the complaints of 460 dogs and cats in the afterlife charging him with cruelty in their deaths.

Coffin Head Li is judged guilty of his crimes and taken away for punishment in the afterlife. This story, like many of the ghost stories of ancient China, served to teach a moral lesson on how one should behave toward others, in this case animals, and the unpleasant fate which awaited one who broke with the kind of behavior expected by the gods.

Treating dogs with respect and remaining true to one's word is one of the morals of the Tale of Panhu. The Chinese ethnic groups of the Yao and She claim the dog Panhu as their ancestor. According to their legends, an emperor named Ku was in conflict with a powerful enemy and promised his daughter in marriage to anyone who could bring him his opponent's head. Panhu the dog accomplished the goal and appeared at the palace with the head and so, however reluctantly, the emperor gave him the princess. The legend concludes with the two of them vanishing into the mountains where Panhu becomes a human prince and he and the princess have many children together.

Dogs were considered so important, they were thought to deserve divine protection for their services.

The transformative nature of the dog is integral to another folktale known as The Lamp Dog. In this story, the steward of a wealthy merchant notices a bright lamp shining above his shelter on his master's property. The light from the lamp seems to slowly descend in a shimmering ball toward the ground where it becomes a dog. The steward rises and follows the dog through the garden where he sees her transform into a beautiful young woman. The steward and the lady spend many nights together until the master hears of this and demands he capture the woman and bring her to him.

The steward does not want to betray his lover but fears his master and so tries to trap her, but she vanishes. Months later, the steward sees his lady as he is coming home from an errand and she leads him to a field of sorghum where a beautiful mansion stands. She tells him their affair is over because of his betrayal but she will give him a farewell party. She prepares him a feast and has many lovely handmaidens wait upon him. When he is done and is returning home, he looks back to see only the field of sorghum.

The lady in the tale is always faithful to the steward, right up until the end when she prepares him the feast, even though he has treated her poorly, just as a dog would behave. The loyalty of the dog and its undeserved mistreatment is at the core of this story as suggested by the woman transforming from a dog.

The loyal dog is featured in many other tales but it is not always seen as a helpful presence. The myth of the Heavenly Dog depicts a dog (known as Tiangou, which literally means 'heavenly dog') which eats the sun or moon during an eclipse and is responsible for childless marriages. Scholar Edward T. C. Werner writes:

The dog is the Dog-star and if the `fate' of the family is under this star there will be no son or the child will be short-lived. Chang Hsien is the patron of child-bearing women and was worshipped by women desirous of offspring. (178)

Chang Hsien is depicted on amulets and portraits from the Sung Dynasty as a bearded man with a raised bow and arrow standing protectively by a young boy. Chang Hsien's job is to shoot the Heavenly Dog before it can cause any trouble. Rituals surrounding Chang Hsien and Tiangou included beating dogs to drive the spirit of Tiangou from a home, and especially so if a couple was trying to conceive a child. Spiritual influence was considered a major factor in conception as, in fact, in every other aspect of life.

Dogs & the Dead

Spirits and the harm they could cause were of great concern to the people of ancient China and remain so today. The belief in ghosts is so prevalent that it even affects the way roads are designed in the country. It is believed that ghosts can only walk in a straight line and so roads are specifically designed to curve and swerve so that certain types of spirits cannot follow one home.

The dog collar also was affected by this belief in ghosts and their manner of travel. Small bells were attached to dog collars, quite common on the collars of the Shih Tzu even today, to scare off spirits. In time, mothers would fasten dog collars to their children with these same kinds of bells to protect them from unseen spirits; a ghost would hear the bells, think the child was a dog, and turn in the other direction. Since a ghost could only walk in a straight line, it was assumed it would not turn back and the child would be safe.

This belief in ghosts stemmed from the understanding that the dead continue to exist after life and are able to help the living (in the form of one's ancestors) or cause harm (as either restless spirits or ones who believe they have been injured or dishonored by the living). Ghosts were potent forces which could easily ruin someone's life if precautions were not taken, and one of the most effective defenses was the dog. Treistman comments on Chinese burial practices and the importance of the dog:

Those who carried the prestige of lineage – and, perhaps at the beginning, those who had acquired special recognition as leaders in raiding and warfare – were buried in large tomb chambers, always accompanied by a dog. (132)

The dog served its master in the afterlife just as it had while on earth by protecting against evil spirits and ghosts. The Chinese afterlife began with the soul crossing a bridge to the other realm where it would either enter heaven to reunite with its ancestors or, if unworthy, drop from the bridge into the abyss of a kind of hell. The dog was thought to go before its master, in some versions of the tale, as both guide and protector. The dog's good nature and loyalty was a comfort on one's final journey although these qualities were also how it almost was excluded as one of the animals of the Chinese zodiac.

The Dog in the Zodiac

The dog is the 11th sign of the Chinese zodiac and among the most popular. According to the story The Great Race, which explains how the zodiac came to be, the Jade Emperor proclaimed there would be a contest and the first twelve animals to cross the finish line would be rewarded with placement in the night sky. Ten animals had already run past the waiting Jade Emperor when the dog finally frolicked by. The Jade Emperor stopped him and asked why he was so late when he always ran so fast. The dog explained that he met friends along the way he had to greet and play with but never lost sight of where he had been called to come.

Another version of the zodiac tale has the dog far off in another country fighting evil spirits and this is what makes him late crossing the finish line. Both versions of the story touch on the nature of the dog as playful, friendly, loyal, and protective. People born under the sign of the dog are considered honest, sincere, loyal, energetic, and intelligent. According to Chinese astrology, their most productive hours are between 7-9 pm (19:00-21:00) and, in the yin-yang dichotomy, they are yang. Their warmth and loyalty attract many friends, and they are usually prosperous and generally happy.


The dog and the long-standing belief in supernatural entities are central to an annual Chinese ceremony known as Tomb Sweeping Day, which falls around the 4th of April during the Qingming Festival. People visit the graves and tombs of their relatives and bring gifts, make sure the graves are cared for, and talk with the dead. Tomb Sweeping Day is not only concerned with the graves of human beings, however, but with one's pets.

The graves of dogs, especially, are honored now during the Qingming Festival, and pet cemeteries have become increasingly popular. From the 1950s through the 1980s it was illegal to even own a dog in Beijing because it was thought a frivolous waste of resources, but the Chinese government has relaxed this stricture, and more and more people in China now own dogs and also spend considerable amounts of money to purchase and maintain their graves once they have passed on.

People visit these cemeteries on Tomb Sweeping Day and leave favorite toys, food, and other objects as presents for their dogs in the same way as for centuries the Chinese have done for their ancestors. The Chinese view of the dog as a resource, rather than companion, would seem to have changed dramatically from ancient times but, as noted, the dog has actually always been more important to the culture than many recognize.

Are These the Oldest Images of Dogs?

Cave art discovered in Saudi Arabia dates back thousands of years and possibly shows hunters leading dogs on leashes.

In northwestern Saudi Arabia, a hunter surrounded by a pack of dogs threads an arrow, props it up against the string of his bow, and pulls it back to kill a wild animal roaming nearby. He's flanked by other hunters readying their weapons.

This scene, and others like it, are engraved onto the cliffsides protruding from the dry, arid desert covering the northern region of present-day Saudi Arabia. A team of researchers think it may be the earliest depiction of dogs ever documented—although other experts aren't yet convinced of that fact. (This story was first reported by David Grimm for Science.)

"The distinction of herders and hunters was immediately clear," said Maria Guagnin, an archaeologist from the Max Planck Institute for the Science of Human History in Germany. Working with the Saudi Commission for Tourism and National Heritage, Guagnin documented 1,405 rock art panels that contain 6,618 individual animal depictions.

Rock art carved into the cliffsides of archaeological sites at Shuwaymis and Jubbah shows images of dogs aiding human hunters. Compared to the adjacent carvings of people, the canines are medium-sized, with short snouts, pointed ears, and a perky, curled tail. They resemble the modern-day Canaan dog breed.

The rock art has an intriguing detail that shows an early effort to use dogs as hunting partners. What appear to be leashes tether the dogs to their human companions. Extending from the dogs' necks, rope-like lines run to human waists. In one scene, the image of a human is shown poised to use a bow to shoot an arrow, with tethered dogs flanking his sides. (Learn why dogs are so friendly.)

China’s history of dog-eating

Dogs have been a part of Chinese households for at least 7,000 years, archaeologists say. The mythological ruler Fu Xi was said to have domesticated six wild animals: the pig, ox, goat, horse, fowl and dog, indicating that dogs were often kept even in ancient times.

Records show that back then dogs were kept mainly to assist with hunting. As the Chinese people became more engaged with agriculture, the dog’s role as hunter became less important – but it was not cast aside. Its loyalty to its owner made it valued for its role as a guard.

Those who advocate the eating of dog maintain it is a Chinese tradition, claiming that historical documents tell of “dog butchers” who specialised in preparing the meat. Others quote from works by founder of the Han dynasty Liu Bang and Qing dynasty painter Zheng Banqiao as proof that the Chinese have always enjoyed dog meat – but this is not enough to prove it is a tradition or custom.

The San Zi Jing, a text used to teach children since the 13 th century, describes dog as one of the six animals raised by people. This is generally taken to mean that these animals were a source of meat. But as agriculture developed and eating habits changed cows, sheep, chickens and pigs became the main sources of meat for Chinese people. Dogs gradually stopped being used as food – and the reasons behind this are complex.

Prior to the Qin and Han dynasties the combination of primitive agricultural techniques and the chaos of constant war meant that living standards were low and meat a rare luxury, offered primarily to the elderly as a sign of respect. Beasts of burden and guard dogs which died of illness or old age could not be wasted, so our thrifty ancestors would cook the meat and eat their fill.

Dog meat was not an essential food for people, as can be seen from a study of sacrificial offerings. These offerings to the gods and their ancestors were important and great attention was paid to the goods to be offered. For the grandest of imperial ceremonies a cow or horse would be sacrificed, for less important occasions a pig or sheep, and the ordinary people would offer pork, chicken or fish. But dog was almost never used, and it was regarded as disrespectful to the spirits to do so. That taboo is still common today, showing that dog meat is not suitable for refined tastes, and certainly not for serving to guests.

Dog fell increasingly out of favour after the Han dynasty. Philosophical Taoism, which rose in the late Han, saw dogs as unclean and consumption of dog was believed to harm efforts to live a simple life. During the Tang and Song dynasties dog consumption decreased further as the range of available meats increased and stories of faithful dogs and Buddhist ideas of reincarnation spread.

China has many ethnic minorities, each with its own traditions and culinary customs. But none of them can be described as dog-eating. In Islam dogs are regarded as unclean and so there is a religious prohibition on eating dog meat. Mongols are traditionally nomads and see dogs as guards and staunch companions. For Manchus eating dog is taboo, due to a legend that a dog saved the life of their forefather Nurhaci. The Tibetans are Buddhist and will not kill animals unnecessarily, and see dogs as loyal companions, so rarely eat them. And even the Zhuang people of Guangxi – where the Yulin “dog-meat festival” takes place – are not recorded in historical documents as being keen dog eaters.

Dynasty Dogs: The Royal History of the Shih Tzu

Did you know that your loyal little Shih Tzu has deep roots in royal Chinese history?

While many associate the breed with Dowager Empress Tzu Hsi, who ruled in China from 1861 through 1908 and considered the dogs sacred, Shih Tzu appear in tapestries dating as far back as 2000 years. While its origins are not fully clear, there is evidence that the breed&mdashdistinct from the Lhasa Apso, Pug, and Pekingese&mdashwas developed by Tibetan Monks who offered the temple dogs as gifts to the emperors of China.

Holy pets of the palace

Called Shih-tzu Kou in traditional Chinese, which literally means &ldquoLion Dog,&rdquo the breed&rsquos lion-like facial features were revered in Imperial courts because Buddha was said to have ridden to earth on the back of a lion.

During the Ming and Manchu Dynasties, the little lion dogs were bred and raised by palace eunuchs and were considered the exclusive property of the royal court. They were rarely seen outside the palaces and anyone caught owning one could be sentenced to death.

These thickly coated Shih Tzus were sometimes carried inside the robes of noble women and were even used as bed warmers and placed at the feet of the emperors and empresses to generate heat.

From China to the West

Dog historians believe that after Empress Tzu Hsi came to power in the 1860s, the Dalai Lama at the time presented her with a breeding pair of extraordinary Shih Tzus. These magnificent dogs were the foundation of her pure line.

Eventually, the dogs were given as gifts to English and Dutch nobility and by 1938 a standard was set for the breed. In the late 1930s, Shih Tzus finally arrived in America and rose to enormous popularity by the 1960s. Even half a world apart from the palaces of China, Shih Tzus are never far away from their royal bloodlines.

Saurr, Norway

Many pet owners are guilty of treating their dogs like royalty, but in the case of Saurr, he truly was a royal dog. The story goes that in the 11th century, King Eystein of Drontheim had conquered the city of Trondheim, which he left to his son Onund to rule.

A whale later, Onund was assassinated. Enraged by this act, King Eystein gave the people a choice: to choose their new king from his slave, Thorer Fax, or his dog, Saurr. The citizens of Trondheim chose the dog, as they believed they’d have their kingdom back and would be free to govern themselves.

Legend has said that this particularly famous dog was gifted with the intelligence and wisdom of three men, and could speak in human tongues. Saurr ruled the kingdom for three years, during which he was treated like royalty, fed only the best food , and was given a gold and silver collar with jewels.

Saurr lived a pampered life as Dog King, but he was also the shepherd to the royal cattle. After ruling the kingdom for three years, Saurr died defending the cattle from a pack of wolves.

Peritas was the famous dog of Alexander the Great and accompanied him during military expeditions. ( Public domain )

Pugs in Tibet

Another theory about the origin of pugs places the breed first in Buddhist monasteries in Tibet around 400 BCE. Monks may have used the pugs primarily as pets and possibly as guard dogs (Pug History). The pug bears a resemblance to Chinese Fu Lions, or Guardian Lions, and it is believed that the sculptors may have modeled the lions after the pug, or one of several other Chinese dog breeds that are similar in appearance (such as the Pekingese). Fu Lion statues stand outside of Chinese temples and are believed to have protective powers. Monks may have kept pugs because of their resemblance to the guardian lions (Sutherland).

In Tibet, pugs were called lags k’yi, which means “hand dog.” This comes from a Tibetan legend about the origins of pugs. According to the legend, if a human touches a young eagle right after it is hatched, it will be transformed into a pug dog (Swainston-Goodger 12).

There is some controversy as to whether pugs were originally bred in monasteries and temples or by the Chinese royalty. One theory claims that pugs were originally bred by Buddhist monks and that their popularity spread to the outsiders before being adopted by Chinese royalty (History of the Pug), but most sources place the breed first as lapdogs and �shion accessories” for royalty before being introduced in Tibet to the Buddhist monks.

According to Tibetan legend, if you touch an eagle right after it hatches, it will be transformed into a pug dog.


Sample collection and whole genome sequencing

58 canids from around the world were gathered for this study. This collection includes 12 gray wolves from across the Eurasian continent, 11 indigenous dogs from southern East Asia, 12 indigenous dogs from northern East Asia, 4 village dogs from Africa (Nigeria) and a set of 19 diverse dog breeds distributed across the Old World and the Americas.

Chinese indigenous dogs are dogs living in the countryside of China 16 (Supplementary information, Data S1 and Figure S1) and were sampled across the geographic range of rural China, including many remote regions in Yunnan and Guizhou in southern China (Supplementary information, Table S1). The breeds include dogs from Central Asia (Afghan Hound) and North Africa (Sloughi), Europe (eight different breeds), the Arctic and Siberia (Greenland dog, Alaska Malamute, Samoyed, Siberian Husky, and East Siberian Laika), the New World (Chihuahua, Mexican and Peruvian naked dog) as well as the Tibetan Plateau (Tibetan Mastiff). These dogs were chosen to cover as many major geographic regions as possible (Figure 1A and Supplementary information, Table S1).

Population structure and genetic diversity of 58 canids. (A) Geographic locations of the 58 canids sequenced in this study. (B) Amount of of SNPs and small indels called in this study. (C) Genetic diversity for the 58 canids. AF, African village dogs BEM, Belgian Malinois CHI, Chihuahua FIL, Finnish Lapphund GAL, Galgo GNE, Gray Norwegian Elkhound GSD, German Shepherd Dog JAM, Jamthund LAH, Lapponian Herder MEN, Mexican naked (hairless) PEN, Peruvian naked (hairless) SWL, Swedish Lapphund AFG, Afghan Hound SLO, Sloughi SAM, Samoyed ESL, East Siberian Laika SIH, Siberian Husky ALM, Alaska Malamute GRD, Greenland dogs TIM, Tibetian Mastiff. (D) Structure analysis of the 58 canids. (E) Genetic diversity of the different groups. AF, African village dogs EB, European breeds SI, southern Chinese indigenous dogs W, wolves. (F) Linkage disequilibrium patterns for the different groups. (G) Principle component analysis of the 58 canids. Inset is for all individuals and the large panel is for dogs only. (H) Principle component plot for a large collection of canids together with our data. (I) A clock-like tree (UPGMA) for all the 58 individuals 56 .

After DNA extraction, individual genomes were sequenced to an average of 15× coverage (Supplementary information, Table S1). Of the 58 individuals, 4 gray wolves and 6 dogs have been sequenced in a previous study 10 . DNA sequence analysis was done using the Genome Analysis Toolkit 17 . After stringent filtering, we identified 20 353 184 SNPs and 3 856 246 small indels (Figure 1B), most of which are shared between groups. For example, 40.3% of the SNPs are shared between wolves, indigenous dogs and dog breeds, reflecting their recent divergence (Figure 1B). Using Sanger sequencing, we verified that the sequencing strategy was highly sensitive (false negative rate around 10%) and the amount of false positives was less than 5% (Supplementary information, Data S2 and Figure S2).

Genetic diversity and population structure

Comparison of the two haploid genomes within each individual yields the genetic diversity θ (4 Nμ) for the 58 individuals. As shown in Figure 1C, genetic diversity shows a decreasing trend from wolves to Chinese indigenous dogs (preserving 78% of the wolf heterozygosity) and subsequently to dog breeds (66% of the wolf heterozygosity), with the African village dogs having a genetic diversity comparable to many dog breeds (69% of the wolf heterozygosity). Among the dog breeds, the levels of variation in genetic diversity are quite dramatic. For example, the East Asian breed Tibetan Mastiff and East Siberian Laika show levels of diversity comparable to the Chinese indigenous dogs, but many of the European dog breeds have considerably reduced genetic diversity. Such dramatic differences in genetic diversity can be influenced both by ancient and recent history of inbreeding.

To explore the genetic relationships among these individuals, we performed a structure analysis using an expectation maximization (EM) algorithm to cluster the individuals into different numbers of groupings 18 . When partitioning the individuals into two groups, the algorithm separates the dogs from the wolves, with very limited admixture observed (Figure 1D). Further dividing the individuals into three subsets split the dogs into two clusters, with indigenous dogs from southern East Asia representing one subset and the other subset consisting of dog breeds from Europe and South/Central America and the African village dogs. Indigenous dogs from northern China and dog breeds from the Arctic and Central Asia, the Middle East and North Africa show a mixture of these components with varying proportions. This observation implies that there are two divergent groups of dogs: one is East Asian component and the other, non-East Asian component. It is important to emphasize that individuals with mixed constituents identified in the structure analysis are not always due to true admixture events, since populations of intermediate genotypes between these two groups tend to display mixed components (e.g., originated shortly after the split of two clades, Supplementary information, Data S3 and Figure S3). Further partitioning into four and five groups leads to the separation of the African village dogs and the breed dogs from the eastern Arctic regions (i.e., Siberian Husky, Alaska Malamute and the Greenland dog).

Genetic diversity among individuals (Figure 1C) may be heavily influenced by ancient as well as recent history, e.g., breeding programs during the last few thousand years or the past few hundred years. However, combined information from multiple breeds may reveal information about the ancestral populations that gave rise to them, since each breed has experienced separate breeding history. We therefore calculated the genetic diversity (θπ) for the “pure groups” informed by the structure analysis (K = 4, Figure 1D). As shown in Figure 1E, dog breeds, most of which of European origin, carry lower diversity than the Chinese indigenous dogs as a group, but have higher genetic diversity than the African indigenous dogs. This suggests that the ancestral population that gave rise to the European breeds was larger than the ancestral population of the African indigenous dogs. Linkage disequilibrium patterns also show similar trends (Figure 1F).

Principle component and phylogenetic analysis

When projecting the genotypes into a two-dimensional space using a principle component analysis (PCA) 19 , all dogs cluster together tightly compared with the distribution seen for wolves (Figure 1G, inset). When inspecting the distribution among dogs, we find that dogs spread along three major geographic axes: southern East Asia, Europe and Africa. The northern Chinese indigenous dogs and dog breeds from the Middle East/Arctic regions/Tibet fall between these three extremes (Figure 1G). The observed pattern reflects the overall geographic locations of these groups following a clear East-West gradient, which matches quite well the observation from our structure analysis.

Combining our dataset with data from a previous SNP array study, which included a larger number of samples 20 , we found that the southern Chinese indigenous dogs together with several East Asian dogs (e.g., Chow Chow, Akita, Chinese Shar-Pei) are closest to wolves (Figure 1H). When the phylogenetic relationships among our 58 samples are inspected, East Asian dogs spread over both sides of the deepest node connecting all dogs, while dogs from other continental areas coalesce into a subclade and then join with East Asian dogs. Thus, East Asian dogs are the most basal lineages connecting to gray wolves (Figure 1I). It is worth pointing out that the genomes of dogs from Oceania (dingoes and New Guinea singing dogs), although being closer to wolves in the PCA plot (Figure 1H), bear strong signals of admixture with gray wolves 6 , which likely reflects their past history of admixture, before they migrated to Australia and New Guinea (Supplementary information, Data S4 and Figure S4).

Admixture analysis

Using the joint allele frequencies among all populations in our study, we infer the split and admixture history among groups of populations using TreeMix 21 . If migration tracks are not allowed, then the relationships inferred from the TreeMix analysis (Figure 2A) directly reflect the patterns observed in our previous analyses including the structure (Figure 1D), the phylogenetic (Figure 1I) and the principal component analyses (Figure 1G). Thus, following the divergence between contemporary wolves and domestic dogs, the first partition within dogs is between the southern Chinese indigenous group and all other dogs. This is then followed by branching of the other dogs, largely matching the geographical distance from southern East Asia: first, dogs from Central Asia, northern China, and eastern Arctic, followed by dogs in Africa, the Middle East, and western Arctic, and the final group including all dog breeds in Europe and South/Central America.

Demographic and migration histories for the domestic dog. (A) Tree topology inferred from TreeMix when no migratory tracts are allowed. The drift parameter is the amount of genetic drift along each population. Further inferred migratory tracts are shown in the bottom-left corner of the panel. The three important nodes are those that we have provided extensive dating information. (B) The PSMC plot for all the individuals. Gray lines plot the benthic δO 18 levels, which are a proxy for global temperature 61 . The span of the current ice age (Quaternary ice age, 2.58M-now) is shown with an arrow. The x-axis is time plotted in log scale and the y-axis is effective population size. (C) Inferred population demographic history between wolves and southern East Asian indigenous dogs using the joint site frequency spectra. (D) A proposed migratory history for domestic dogs across the world based on the evidence from our study. Solid arrows represent migratory tracts that we have dating information, while dashed arrows indicate those without accurate dating.

If migration tracks are allowed in TreeMix, there is strong statistical support for migrations among a few groups: (1) northern Chinese indigenous dogs show strong admixture from European dogs (Figure 2A and Supplementary information, Data S5, Figure S5, Tables S2 and S3) (2) gene flow from wolves to the African/Middle Eastern dogs (Supplementary information, Figure S5) (3) migratory tracks from the southern Chinese dogs to the eastern Arctic group (i.e., Siberian Husky, Alaska Malamute and the Greenland dog Supplementary information, Figure S5). When all possible migration events in the history of these samples are examined using the F3/F4 test 22 , there is again a strong statistical support for all the migration events listed above (Supplementary information, Data S5).

Long-term evolutionary trajectories for wolves and dogs

Using the divergence between the two haploid genomes within individuals, the pairwise sequentially Markovian coalescent (PSMC) model provides a method for investigating the long-term trajectories in population sizes 23 . To translate demographic history into real-time units, estimation of an accurate mutation rate is very important. Previously, several different mutation rates were used, but they were generally not carefully calibrated (Supplementary information, Data S6) 24 . Using multiple outgroup species to the dog (e.g., horse and cat), our estimate of the mutation rate for the lineage leading to the domestic dog is 2.2 × 10 −9 per site per year (Supplementary information, Data S6 and Table S4), a rate similar to those from several earlier studies 25,26 . Using this mutation rate, we estimate dates for the population history of dogs and wolves. As shown in Figure 2B, a decrease in the size of the ancestral wolf population started to occur 2 million years ago, reaching a saddle point about 3-400 000 years ago. The ancestral population then increased in size, peaking at around 200 000 years ago. After a subsequent small decline in population size, wolves and dogs started to diverge from each other between 20 000 and 100 000 years ago (see next section for a more precise dating). Although all domestic dogs drastically decreased in population size after the population split, the wolf population experienced a slight growth, possibly as a consequence of the megafauna extinctions (i.e., late Quaternary extinction) 27 that provided gray wolves with better food resources due to reduced competition from other predators.

Time of divergence between contemporary wolves and dogs

Treemix and phylogenetic analyses identified southern Chinese indigenous dogs as the most basal population compared to wolves, from which all other dog populations diverged. We therefore used joint allele frequencies between the 12 gray wolves and the 11 southern Chinese indigenous dogs, to infer the demographic history for these two populations with the dadi package 28 . Similar to the result from the PSMC analysis, the wolf population experienced a very mild population growth (1.26-fold increase) that started around 290 000 years ago (Figure 2C). The time of divergence for the wolf and dog populations is inferred to be around 33 000 years ago, where the domestic dog lineage expanded from a population of 4 600 individuals to about 17 500.

In addition to gauging changes in population size, statistical methods can also estimate the rates of exchange of migrants between two populations. The migration rate (2Nm) from the dog lineage to the wolf lineage is estimated to be 0.97, while the other direction (wolves to dogs) is inferred to be 5.02, showing a clear asymmetry in the migration rates 29 .

Examination of the sequence divergences between the multiple populations using a Markov chain Monte Carlo (MCMC) approach 30,31 (Supplementary information, Data S7, Figures S6-S8, Tables S5 and S6) reveals a similar profile for the history between wolves and dogs, which includes a slight growth in the wolf population and an ancient divergence between wolves and dogs (Supplementary information, Data S7 and Table S5). In summary, multiple levels of genetic information (i.e., both joint site frequencies as well as sequence divergence) support an ancient split between dogs and wolves.

The geographical origins of dogs: a single origin in southern East Asia

In order to identify the most probable geographical origin of dogs, we hypothesized that similar to many organisms, the geographical origin of a species holds the greatest genetic diversity, and the global relationship among multiple populations will, in the absence of strong influence of admixture, follow a serial founder model 32,33 . In the case of dogs, the wild ancestor, the wolf, has been present along the dog throughout Eurasia, implying that intense dog-wolf admixture could possibly have influenced this pattern.

Despite the concern on the confounding effect of wolf/dog gene flow, the TreeMix analysis, F3/F4 test as well as the demographic analysis suggest that gene flow between dogs and wolves is relatively mild. In Supplementary information, Data S8, we review the evidence for dog/wolf gene flow from our study, as well as from multiple previous studies. The combined evidence shows that the migration rates (2Nm) are mostly around one or less (a maximum of five found in the dadi analysis) and that the admixture proportion is normally around 10%, with a maximum of 16% for the Middle East (Supplementary information, Data S8). Low levels of migration are detected between wolves and dogs across Eurasia when the very sensitive D test is used 34,35 (Supplementary information, Data S8). Thus, we conclude that while dog-wolf gene flow has occurred throughout history of the domestic dog, it has been at a moderate level and the level of admixture has been relatively similar across Eurasia (Supplementary information, Data S8). Without the strong influence of admixture 32 , we may assume that genetic diversity is highest at the place of origin and that the global relationship among the multiple populations follows a serial founder model reflecting their dispersal routes 33 .

It is tempting to draw conclusions about the origin of dogs from the high genetic diversity observed in the Chinese indigenous dogs. However, comparing breed dogs with indigenous dogs at the individual level is likely misleading since most of the differences in genetic diversity are probably caused by recent bottleneck events rather than their distant origin 1 . Thus, we combine multiple breeds in each region as a group representing the ancestral haplotype pool giving rise to the contemporary dogs of that region. Our analysis shows that dogs from East Asia have the highest genetic diversity (Figure 1E). This suggests that the ancestral population that gave rise to East Asian dogs was much larger than ancestral populations in other regions (e.g., Europe). The linkage disequilibrium pattern also shows the same trend (Figure 1F). Higher levels of genetic diversity in East Asian dogs are also observed in mtDNA and Y chromosome data 7,12,36 .

Beside group diversity, in the phylogenetic and TreeMix analyses, the deepest node connecting all dogs separates into two clades, one of which is composed of only East Asian dogs, while the other clade includes both East Asian and non-East Asian dogs (Figures 1I and 2A, and Supplementary information, Figure S5). Dogs from Africa and Europe share a most recent common ancestor, which then coalesces with dogs from East Asia (Figures 1I and 2A). Notably, this basal position of East Asia is robust to the levels of migrations between wolves and dogs (Supplementary information, Data S9, Figure S9, and Table S7). The basal position of East Asian dogs is similar to the pattern observed for Africans within human populations 37 .

In addition to the observations based on group level diversity and the basal phylogenetic position, the PCA pattern also provides supporting evidence for the southern East Asian origin of dogs. As the amount of genetic drift in basal groups is typically lower due to their larger population sizes, we expect them to display a closer genetic relationship with wolves in the PCA plot (Figure 2A). When we simulate a serial founder model that mimics the history of dog domestication, we can easily generate a pattern that is similar to that shown in Figure 1G (see also Supplementary information, Data S10 and Figure S10). Thus, in our analysis, we find dogs with ancestry in southern East Asia to be closest to wolves, and also a geographical distribution of the populations following a clear east-west gradient, indicating serial founder events. It is important to emphasize that admixture between wolves and dogs is unlikely to have created the observed pattern, given that the dog-wolf admixture rate in East Asia is not higher than that seen in other regions (Supplementary information, Data S8).

Having identified southern East Asia as the likely origin of dogs, we asked whether the domestic dog may have originated in more than one region through separate domestication events. In order to test whether multiple origins are compatible with the observed data, we performed simulations mimicking different scenarios (Supplementary information, Data S11 and Figure S11). Our results show that, if there were multiple origins for dogs from separate wolf populations, the descendant populations would tend to reside in separate clusters in the PCA plot, which is in contrast to what we observe (Figure 1G, inset). Thus, that the domestic dog originated multiple times in different geographical areas is not compatible with the observed genetic patterns found in our genome data.

The out of southern East Asia history for the domestic dog

To study the subsequent global history of the dog, we used an MCMC approach to date several important transitional points among the major clades (Figure 2A). Our analysis supports the split between the southern Chinese indigenous dogs and all other dogs across the world around 15 000 years ago, thus indicating a radiation of dogs out of southern East Asia earlier than the origin of agriculture (Supplementary information, Data S7 and node 2 in Figure 2A and 2D) 38 . After radiating from southern East Asia, possibly following existing human settlements at the time (Supplementary information, Data S12 and Figure S12), the out of southern East Asia lineage spread to the Middle East/Africa and arrived in Europe by about 10 000 years ago (Supplementary information, Data S7 node 3 in Figure 2A and 2D). Notably, one of the out of southern East Asia lineages migrated back to northern China, meeting endemic Asian lineages that had spread from southern East Asia and yielding a series of admixed populations, including the northern Chinese indigenous dogs and the Arctic dog breeds (Figure 2A and 2D).

Several dog breeds from South and Central America (i.e., Chihuahua, the Mexican and Peruvian naked dog) show no signs of admixture, while the Arctic breeds, Alaska Malamute and the Greenland dog, display extensive admixture from the southern Chinese Indigenous lineage 39 . Possibly, this reflects that the human colonization of the New World occurred in several waves, in which dogs may have followed in different time periods 40 (Figure 2D). Using the patterns of the admixture tracks, we estimate that the time of the admixture for the northern Chinese indigenous dogs was quite ancient (around 10 500 years ago, Supplementary information, Data S13 and Figure S13) 40 . The relatively recent origin of European dogs (i.e., ∼ 10 000 years) together with this rather ancient admixture suggests that multiple lineages travelled to the Far East from the Middle East/Europe.

Population structure among wolves

Our structure and principal component analyses do not reveal any population substructure among the gray wolves collected for this study (Figure 1D). The high migratory ability of the gray wolf might allow the populations to remain highly homogenous across the eastern part of Eurasia 41 . A previous study using wolves from the Middle East (Israel), Europe (Croatia) as well as China found genetic differentiation among these wolf populations 6 . When these three individuals are overlaid on the large PCA plot, the wolves from western Eurasia do not group together with the wolves we collected from eastern Eurasia, and they are genetically closer to dogs (Supplementary information, Data S14 and Figure S14). Given the fact that Middle Eastern wolves generally have more dog admixture 6 , the observed difference might not represent true population differentiation among wolves. Nevertheless, it is possible that some wolves have recently diverged from each other 8 , as there is weak isolation between the wolves from eastern and western Eurasia. Explicit testing for potential admixture between wolves and dogs sampled in our study finds evidence of gene flow between wolves and local dog populations in each region, albeit the magnitude is low (Supplementary information, Table S8). Further study on the genetic and geographic relationships between dogs and wolves is one of the important tasks for the community.

Domestication genes

Our analyses indicate that the Chinese indigenous dogs represent an intermediate form between wolves and breed dogs, and they have not experienced intense artificial selection. Analyses of Chinese indigenous dogs therefore allow us to stratify the domestication process in dogs, and investigate the role of positive selection that occurred specifically during the first stage of domestication. Using a statistical method that explicitly models selective sweeps 42 , we have identified the top 1% of the genome bearing strong statistical evidence of positive selection in the southern Chinese indigenous dogs. In Table 1, we list the categories of genes that show statistical significance by a gene enrichment-based analysis. Groups of genes showing the strongest evidence of positive selection are those related to metabolism and motility, neurological process and perception as well as sexual reproduction (Table 1 and Supplementary information, Data S15, Tables S9 and S10). Genes that seem to have been positively selected in subsequent evolutionary steps, including dog breed formation, are related to the control of developmental processes and to metabolism (see a full discussion of candidate genes involved in transforming wild wolves to dogs in Supplementary information, Data S15).

Among the candidates as positively selected genes in the first stage of dog domestication, a class of genes are related to memory and long-term potentiation (LTP), which is widely considered to be the major cellular mechanism underling learning and memory 43 . For example, GRIA1 (glutamate receptor, ionotropic, AMPA 1) is an important protein that mediates excitatory synaptic transmission in the central nervous system and plays a key role in hippocampal synaptic LTP and long-term depression (LTD). Interestingly, a suite of other genes, including GRIN2A (glutamate receptor, ionotropic, N-methyl D-aspartate 2A), are also found to be heavily involved in LTP and LTD (Table 1). The large physiological and behavioral changes empowered by these genes may have enabled the transformation of gray wolves to domestic dogs, allowing them to flourish in the human environment.

Chinese Dogs: 9 Awesome Dog Breeds From China

They can be poofy. They can be pug-faced. They can be distinct. And more often than not, Chinese dogs are incredibly cute. Chinese dogs also generally have long, storied histories. Not all the breeds considered to be Chinese actually originate in China.

If you're looking for a truly Chinese dog breed, here are nine distinct dogs to consider. The best way to go about adopting a Chinese dog is to find a registered breeder at the American Kennel Club website.

    The Pekingese has long been associated with the history of China and its emperors. Records of the Pekingese (named after Peking, the former name for Beijing, the capital of China) date back to at least 700 C.E. Its long, regal coat requires about an hour per week of brushing. But the Pekingese is low-maintenance in personality and generally known as a chill, indoor-type dog. According to the American Kennel Club, the introduction of the Pekingese to the Western world stemmed from the looting of the Imperial Palace in Beijing in 1860.

Erin Chan Ding is a freelance journalist in the Chicago area who writes about travel, pets, news, business, parenting and features.

Domestic Ancient China Animals have been used in China even before the Tang dynasty. The Tang dynasty paintings and sculptures depicted domestic animals in various forms, making them an integral part of ancient Chinese culture.

Dogs in ancient China

Dogs were a part of human settlements and households even in the prehistoric times, though the breed is not known.

During the Shang dynasty, the construction of every palace, tomb or royal building was concluded only with the help of a dog sacrifice. Dogs were sacrificial victims even in the Ning rites when they were dismembered into four parts as an act of worship of the four directions.

During the Chou dynasty, too dog sacrifices continued to hold great significance. The ba sacrifice was regarded as an act to ward off evil and was carried out in the presence of the Emperor himself.

Dogs were also a source of protein in ancient China and dried fish fried in dog fat was a regular preparation to reduce body heat during summers. Even the Emperor had to eat dog meat during the first three months of autumn as it was considered to reduce fatigue.

Horses in Ancient China

Horses find a mention in even during the Shang dynasty when horse-drawn chariots were already in use. It was in 4th century BC that people started riding on horsebacks. Horses were used for hunting and warfare and though they were also used for sacrifices, the practice died out as horses became very expensive.

Ancient Chinese Zodiac Animals

The Chinese zodiac also comprises of animals associated with each year. They include the rat, pig, dog, rooster, monkey, sheep, horse, snake, dragon, rabbit, tiger and the ox. It is said that the zodiac originally did not consist of any animals and that Lord Buddha later assigned these animals for every year.

Mythical Animals of Ancient China

China has a rich and significant tradition of worshipping mythical animals like the unicorn, the famous dragon, and the Phoenix.

The unicorn is worshipped for long life, contentment, prosperity, and blessings for the offsprings. The Phoenix is a symbol of the sun, agricultural wealth, and good harvest.

The dragon is the most famous Chinese symbol and since ancient times is regarded as an animal with divine powers. It is also considered as a symbol of culture.

Other Animals in Ancient China

The tortoise is yet another animal that is greatly revered in China since ancient times. It is considered as a symbol of strength and endurance.
The other animals that find a mention in ancient Chinese history include the panda, the leopard, and the monkey.

A glimpse of the animal life in ancient China indicates the great significance that people attached to animals in daily life.

Science Shines More Light on Cats in Chinese History

More details about the domestication of cats in ancient China emerged in January of 2016. A new team of scientists headed by French researchers at the CNRS to identify the species of cat that had been found by the National Academy of Sciences. They wanted to know if the cats were of the same species and what relationship this species had with the Chinese people.

Since DNA evidence was not viable, geometric metamorphic analysis was used on the bones. The results were somewhat surprising. It was determined that the bones were those of a common east Asian wildcat known as the Leopard Cat (Prionailurus Bengalensis). These are the wildcats that were used to help create the modern hybrid cat breed, the Bengal.

What was the surprise? The Leopard Cat is not the ancestor of the domestic cat (Felis Catus) known today. Rather, the domestic cat is descendant from a distant relative of Leopard Cat, the African Wildcat (Felis silvestris lybica). There is no record of the African Wildcat living in China despite all of the current domestic cats being of the Felis Catus species.

This new revelation leaves scientists with even more questions about cats in Chinese history. The domestication of cats had independently cropped up in 3 separate locations in the world (Egypt, the Middle East, and China) as each began to develop agriculture. How and when did the domestic cat replace the Leopard Cat in the process of domestication in China? Was it the Romans, trade routes with the west, or something else that brought domestic cats to China? Only further studies into cats in Chinese history will tell.

Watch the video: Secrets Of The Great Wall. Ancient China From Above. National Geographic