Page semi-protected

Origin of the domestic dog

From Wikipedia, the free encyclopedia
Jump to: navigation, search
This article is about the origin of the domestic dog. For dog breeding, see Dog breeding.

The origin of the domestic dog (Canis lupus familiaris or Canis familiaris) is not clear. Whole genome sequencing indicates that the dog, the gray wolf and the extinct Taymyr wolf diverged at around the same time 27,000-40,000 years ago.[1] These dates imply that the earliest dogs arose in the time of human hunter-gatherers and not agriculturists.[2] Modern dogs are more closely related to ancient wolf fossils that have been found in Europe than they are to modern gray wolves,[3] with nearly all genetic commonalities with the gray wolf due to admixture [2] but several Arctic dog breeds with the Taymyr wolf of North Asia due to admixture.[1]

The dog diverged immediately prior to the Last Glacial Maximum when much of Eurasia was a steppe/tundra biome


The dog came into being towards the last glacial maximum in very cold and dry climatic conditions.

The last glacial period, commonly referred to as the 'Ice Age', spanned 125,000[4] to 14,500[5] years ago and was the most recent glacial period within the current ice age which occurred during the last years of the Pleistocene era.[4] The Ice Age reached its peak during the last glacial maximum (LGM), when ice sheets commenced advancing from 33,000 years BP and reached their maximum positions 26,500 years BP. Deglaciation commenced in the Northern Hemisphere approximately 19,000 years BP, and in Antarctica approximately 14,500 years BP which is consistent with evidence that this was the primary source for an abrupt rise in the sea level 14,500 years ago.[6]

A vast Mammoth steppe stretched from Spain across Eurasia and over the Bering land bridge into Alaska and the Yukon where it was stopped by the Wisconsin glaciation. This land bridge existed because more of the planet's water was locked up in glaciation than now and therefore the sea-levels were lower. When the sea levels began to rise this bridge was inundated around 11,000 years BP.[7] During the last glacial maximum, the continent of Europe was much colder and drier than it is today, with polar desert in the north and the remainder steppe or tundra. Forest and woodland was almost non-existent, except for isolated pockets in the mountain ranges of southern Europe.[8]

The fossil evidence from many continents points to the extinction mainly of large animals at or near the end of the last glaciation. These animals have been termed Pleistocene megafauna. The most common definition of megafauna is an animal with an adult body weight of over 44 kg. Across Eurasia, the Straight-tusked elephant became extinct between 100,000–50,000 years BP. The hippopotamus, interglacial rhinoceros (Stephanorhinus), cave bear (Ursus spelaeus), and heavy-bodied Asian antelope (Spirocerus) died out between 50,000-16,000 years BP. The spotted hyena, woolly rhinoceros and mammoths died out between 16,000-11,500 years BP. The musk ox died out after 11,500 BP, as did the giant deer (Megaloceros) with the last pocket having survived until about 7,700 years BP in western Siberia.[9] A pocket of mammoths survived on Wrangel Island until 4,500 years BP.[10] As some species vanished, so did their predators. Among the top predators, the sabre-toothed cat (Homotherium) died out 28,000 years BP,[11] the cave lion 11,900 years BP,[12] and the leopard in Europe died out 27,000 years BP.[13] The Late Pleistocene was characterized by a series of severe and rapid climate oscillations with regional temperature changes of up to 16°C, which has been correlated with megafaunal extinctions. There is no evidence of megafaunal extinctions at the height of the LGM), indicating that increasing cold and glaciation were not factors. Multiple events appear to also involve the rapid replacement of one species by one within the same genus, or one population by another within the same species, across a broad area.[14]

Modern human's ancestors first appeared in East Africa 195,000 years ago.[15] Some migrated out of Africa 60,000 years ago with one group reaching Central Asia 50,000 years ago.[16] From there they reached Europe, with human remains dated 43,000-45,000 years BP discovered in Italy,[17] Britain,[18] and in the European Russian Arctic 40,000 years ago.[19][20] Another group left Central Asia and reached the Yana River, Siberia, well above the Arctic circle 27,000 years ago.[21] Modern humans then made their way across the Bering land bridge and into North America between 20,000-11,000 years ago, after the Wisconsin glaciation had retreated but before the Bering land bridge became inundated by the sea.[22] These people then populated the Americas. In the Fertile crescent the first agriculture was developing 11,500 years ago.[23]

Into this environment came the dog.

Wolf-like lineage

Early DNA studies indicated that the dog is descended from a wolf-like lineage.

In 1868, Charles Darwin proposed that domestic dogs were phenotypically so diverse that they likely had originated from 2 or more wild canis species.[24]:16 All species within the Canis genus, the wolf-like canids, are phylogenetically closely related with 78 chromosomes and can potentially interbreed.[25] Later, others thought that "the wolf is the most probable ancestor and closest relative of the domestic dog."[26]:54

The development of molecular biology allows the inference of evolutionary relationships about species and to represent them in a phylogenetic tree, however these are not without their limitations. DNA studies may give unresolvable results due to the specimens select, the genome technology used, and the assumptions made by the researchers.[27] A panel of Genetic markers can be chosen from e.g. mitochondrial Cytochrome b. The techniques used to extract, locate and compare sequences can be applied using advances in genome technology to observe longer lengths of base pairs that give better phylogenetic resolution.[28] These techniques can be applied to the maternal mitochondrial control region, mitochondrial D-loop, mitochondrial genome, paternal Y-chromosomes, microsatellites, Single nucleotide polymorphisms, nuclear DNA and the whole genome.

In June 1993, a study of 736 base pairs of the mitochondrial Cytochrome b gene of the wolf-like canids found that there was a close kinship between domestic dogs, gray wolves, coyotes and Simien jackals but with a distance from the African wild dog and from the golden, side-striped and black-backed jackals. The domestic dog was "an extremely close relative of the gray wolf, differing from it by at most 0.2% of Mitochondrial DNA [mDNA] sequence. In comparison, the gray wolf differs from its closest wild relative, the coyote, by about 4% of mDNA sequence." Therefore, the study concluded that the molecular genetic evidence did not support theories that dogs arose from jackal ancestors. The study warned against "the need for caution in the interpretation of phylogenies based on mDNA; such gene trees are not necessarily species trees and may not accurately reflect the phylogenetic affiliations or divergence time." The study proposed the hypothesis that because of the diversity of dog remains found in archaeological sites, that dogs may be derived from several different ancestral gray wolf populations.[29]

In the same year 1993, the domestic dog Canis familiaris was reclassified as Canis lupus familiaris, a subspecies of the gray wolf Canis lupus, in Mammal Species of the World.[30] However, Canis familiaris is also accepted due to a nomenclature debate over the naming of wild and domestic sub-species.[31]

In 1995, a researcher found that the archaeological record did not show any distinctive dog breeds until 3,000-4,000 years ago and that most breeds had been developed over the past seven hundred years.[32]

In 1997, a study of 261 base pairs on the control region of 140 dogs across 67 breeds and 162 wolves across 27 populations. The control region of the dogs and wolves was highly polymorphic, with dogs revealing 26 haplotypes, and wolves revealing 27 haplotypes of which 4 had a widespread distribution. Dog haplotypes could not be partitioned into breeds e.g. 8 German Shepherds revealed 5 distinct sequences, and 6 Golden Retrievers revealed 4 sequences. No dog sequence differed from any wolf sequence by more than 12 substitutions but differed from coyote sequences by at least 20 substitutions and 2 insertions, clearly supporting a wolf ancestry for dogs. The 26 dog haplogroups formed 4 distinct clades; Clade 1 included 19 haplotypes, Clade 2 included 1 haplotype, Clade 3 included 3 haplotypes, and Clade 4 included 3 haplotypes which were identical or very similar to a wolf haplotype found in Rumania and Western Russia and which suggests recent hybridization. A further analysis of 1,030 base pairs on the control region of 24 dogs provided greater phylogentic resolution and supported 4 monophyletic clades. The wolf and coyote ancestral lines diverged one million years ago based on the fossil record, and comparing their sequence divergence with those of the dog implies that dogs diverged 135,000 years ago. The estimate may be inflated as it is based on some assumptions, however it implies that dogs origin is more ancient than 14,000 YBP suggested by the archaeological record. The study proposed the hypothesis that because dog haplotypes form 4 monophyletic clades, that early domestic dogs may not have been morphologically distinct from their wild relatives until the change 10,000-15,000 years ago when humans moved from nomadic hunter-gatherer societies into agricultural societies and imposed selective regimes that resulted in marked phenotopic divergence from wolves. After the origin of dogs from a wolf ancestor, dogs and wolves may have continued to exchange genes.[33]

In 1999, a review of the scientific literature regarding the genetic origin of the dog proposed a number of hypotheses. The molecular data indicated that dogs have protein alleles in common with wolves, share highly polymorphic microsatellites, and have mitochondrial DNA sequences similar or identical to those found in gray wolves. The mitochondrial control region DNA sequences shows an average divergence between dogs and wolves at 1.5% compared to dogs and coyotes, their next-closest relative, at 7.5%. Therefore, this indicated the origin of the dog was from wolves. The archaeological record suggests that dogs were in Europe and the Middle East approximately 14,000 years ago but the genetic record shows 135,000 YBP, which indicates that the morphological change was associated with artificial selection as humans shifted from hunter-gatherer to agrarian societies. Alternately, dogs may have had a more recent origin but are descended from a now extinct species of canid whose closest living relative was the gray wolf. The DNA sequences of dogs form 4 distinct clades, each with a separate ancestry from wolves that indicates 4 separate domestication events. One of the clades shows a wolf sequence that is identical to a dog sequence, suggesting a very recent interbreeding or domestication event. Once dogs were domesticated and spread over a wide area, occasional interbreeding would have transferred wolf mDNA to them.[25]

Probable ancestor

During the LGM, there were two types of wolf. A large, heavily-built megafaunal wolf spanned the cold north of the holarctic that specialised in predating on megafauna. Another more gracile form lived in the warmer south in refuges from the glaciation. When the planet warmed and the LGM came to an end, whole species of megafauna became extinct along with their predators, leaving the more gracile form to dominate the holarctic. This wolf we know today as the modern gray wolf, which is the dog's sister but not its ancestor - the dog shows a closer genetic relationship with the extinct megafaunal wolf.

In 2010, a study compared the mDNA haplotypes of 947 modern gray wolves from across Europe with the published sequences of 24 Pleistocene wolves from western Europe dated between 1,200-44,000 years BP. The study found that phylogenetically the haplotypes represented two haplogroups and referred to these as haplogroup 1 and 2. The 947 European wolves revealed 27 different haplotypes with haplogroup 1 forming a monophyletic clade, and all other haplotypes forming haplogroup 2. Comparison with gray wolves from other regions revealed that haplogroups 1 and 2 could be found spread across Eurasia but only haplogroup 1 could be found in North America. The Pleistocene wolf samples from western Europe all belonged to haplogroup 2, which suggested a long-term predominance in this region. A comparison of current and past frequencies indicated that in Europe haplogroup 2 became outnumbered by haplogroup 1 but in North America haplogroup 2 became extinct and was replaced by haplogroup 1 after the Last Glacial Maximum.[34] Access into North America was available between 20,000-11,000 years ago, after the Wisconsin glaciation had retreated but before the Bering land bridge became inundated by the sea.[35] Therefore, haplogroup 1 was able to enter into North America during this period.

Analysis of stable isotopes, which offer conclusions about the diet and therefore the ecology of the extinct wolf populations suggest that the Pleistocene wolves from haplogroup 2 mainly preyed on Pleistocene megafaunal species,[36][37] which became rare at the beginning of the Holocene 12,000 years ago.[38]:2 "Thus, Pleistocene wolves across Northern Eurasia and America may actually have represented a continuous and almost panmictic population that was genetically and probably also ecologically distinct from the wolves living in this area today."[39]:R610 "The Pleistocene Eurasian wolves are morphologically and genetically comparable to the Pleistocene eastern-Beringian wolves."[40]:791 Some of the ancient European and Beringian wolves shared a common haplotype (a17).[34]:8 The specialized Pleistocene wolves, thus, did not contribute to the genetic diversity of modern wolves. Rather, modern wolf populations across the Holarctic are likely be the descendants of wolves from populations that came from more southern refuges as suggested previously[41] for the North American wolves.[39]:R611

These 2 haplogroups exclude the older-lineage Himalayan wolf and the Indian gray wolf.

See also: Beringian wolf
See also: Megafaunal wolf

The fossil remains of the direct ancestor of the dog have yet to be found and so the probable ancestor is not yet confirmed.

First divergence

Time of divergence

The ancestral dog and the ancestral modern gray wolf diverged from a common ancestor at least 27,000 years ago
DNA evidence indicates that the dog, the modern gray wolf (above) and the now-extinct Taimyr wolf triverged from an extinct wolf-like canid that lived in Europe.

In May 2015, a study was conducted on a partial rib-bone of a wolf (named Taimyr-1) found near the Bolshaya Balakhnaya River in the Taymyr Peninsula, North Asia, that was AMS radiocarbon dated to 34,900 years BP. The sample provided the first draft of the entire nuclear genome of a Pleistocene carnivore and the sequence was deposited in the European Nucleotide Archive (ENA) with accession number PRJEB7788 and classified as Canis lupus because the genome sequence was found to be substantially closer to modern gray wolves than it was to modern coyotes. The data was compared to the genotypes of 532 dogs from 48 breeds and 15 gray wolves from Europe, the Middle East, China, and North America.[1]

Using the Taimyr-1 specimen's radiocarbon date, its genome sequence and that of a modern wolf, a direct estimate of the genome-wide mutation rate in dogs/wolves could be made to calculate the time of divergence. The data showed that the Taimyr-1 lineage was separate to modern wolves and dogs and indicated that the Taimyr-1 genotype, gray wolves and dogs triverged from a now-extinct common ancestor before the peak of the Last Glacial Maximum 27,000-40,000 years ago. The separation of the dog and wolf did not have to coincide with selective breeding by humans.[1]:page3[42]

This derived mutation rate was much slower than that assumed in previous studies. Such an early divergence is consistent with several paleontological reports of dog-like canids up to 36,000 years old, as well as evidence that domesticated dogs most likely accompanied early colonizers into the Americas.[1]

The ancestral fossils have not yet been found.

Place of divergence

Modern dogs show a closer genetic association with ancient, extinct canids from Europe[3] and Arctic north-east Siberia.[43]

A number of mitochondrial DNA studies have searched for the time and place that the dog first diverged from the extant gray wolf, including 135,000 years ago,[44] in East Asia 15,000 years ago,[45] in Africa,[46] in southeastern Asia south of Yangtze River 5,400–16,300 years ago,[47] in the Middle East,[48] in East Asia south of the Yangtze River,[49] and in Europe.[50] Genetic studies comparing the dog to the extant gray wolf did not result in agreement among researchers.

In 1934, an eminent paleontologist indicated that the ancestor of the dog lineage may have been the extinct Canis lupus variabilis.[51]

The 14,500 year old upper-right jaw found in Kesslerloch Cave, Switzerland is the sister to 2/3 of modern dogs (courtesy Hannes Napierala)


In November 2013, a study analysed the complete and partial mitochondrial genome sequences of 18 fossil canids dating from 1,000 to 36,000 years ago from the Old and New Worlds, and compared these with the complete mitochondrial genome sequences from 49 modern wolves and 77 modern dogs—including divergent dog breeds, such as the Basenji and dingo—3 Chinese indigenous dogs, and 4 coyotes totaling 148 mitochondrial genomes. Mitochondrial DNA was analyzed because there is a higher abundance of mitochondrial than nuclear DNA available from ancient specimens. The data indicate that 22% (17 of 77) of the dogs sampled are sister to modern wolves from Sweden and the Ukraine with a most recent common ancestor 9,200 years ago (else admixture with wolves as dogs were clearly domesticated by this time), and 78% (60 of 77) are sister to one or more ancient canids from Europe. Some 12% (9 of 77) of the dogs are sister to two morphologically distinct ancient dogs from Germany, one 14,700 years old from Bonn-Oberkassel (GenBank accession number KF661093) and one 12,500 years old from the Kartstein cave (KF661094) with a most recent common ancestor 16,000–24,000 years ago. [Note: the study's Figure 1 shows the Phylogenetic relationships and indicates that this most recent common ancestor was also dog-like, and separated from what was to become the ancestor of many of the modern Eurasian wolf lines at 24,000–31,000 years ago.] Some 3% (2 of 77) of the dogs are sister to the 14,500 year old wolf sequence from the Kesslerloch cave in Switzerland (KF661087) with a most recent common ancestor 18,300 years ago. They are also distantly rooted in the same sequence as the Altai dog, however the study does not support its recent common ancestry with most modern dogs. Some 64% (49 of 77) of the dogs are sister to another 14,500-year-old wolf sequence also from the Kesslerloch cave in Switzerland (KF661091) with a most recent common ancestor 32,100 years ago. This group of dogs matches 3 fossil pre-Columbian New World dogs between 1,000 and 8,500 years old. Matching these 3 to the 49 relatives indicates a most recent common ancestor 18,800 years ago, which supports the hypothesis that pre-Columbian dogs in the New World share ancestry with modern dogs. Thus, these dogs likely arrived with the first humans to the New World. The early dog population appears to have undergone a population bottle-neck (decrease) between 5,000 and 2,000 years ago, followed by a sharp increase that parallels the trajectory of the human population.[3]

The study sequenced 3 haplotypes from the ancient Belgium canids (the Goyet dog - Belgium 36,000 years BP cataloged as Canis species KF661079, and with Belgium 30,000 years BP KF661080 and 26,000 years BP KF661078 cataloged as Canis lupus) and found they formed the most diverging group. This degree of divergence implied a European origin.[3]

The data from this study indicate an association of modern dogs with ancient European canids and some with modern European wolves, with no close affinity to modern wolves from the Middle East or East Asia. This suggests the origin of dogs from Europe, rather than the Middle East or East Asia, as previously suggested. Additionally, there are no ancient dog remains from these regions older than 13,000 years. Divergence times implies a European origin of the domestic dog dating 18,800–32,100 years ago, which supports the hypothesis that dog domestication preceded the emergence of agriculture and occurred in the context of European hunter-gatherer cultures. An evolutionary scenario consistent with these results is that dog domestication was initiated close to the Last Glacial Maximum when hunter-gatherers preyed on megafauna.[3]

Arctic North-East Siberia

In 2015, a study looked at the mitogenome contol region sequences of 13 ancient canid remains and one modern wolf from 5 sites across Arctic north-east Siberia. The 14 canids revealed 9 haplotypes, 3 of which were on record and the others unique. Four of the Siberian canids dated 28,000 YBP, and one Canis c.f. variabilis dated 360,000 YBP, were as divergent as the ancient European specimens found in an earlier study, and therefore a European origin of domestic dogs may not be conclusive.[43]

The phylogenetic relationship of the extracted sequences and other sequences available from Genbank showed the haplotype from specimen S805 (28,000 YBP) was one step away from another haplotype S902 (8,000 YBP) that represents one of the main phylogenetic clades (Clade A) that include most of the domestic dog lineages. Several ancient haplotypes were oriented around S805 and these include Canis c.f. variabilis (360,000 YBP), Belgium (36,000 YBP - the "Goyet dog") and Belgium (30,000 YBP), and Konsteki, Russia (22,000 YBP). The ancient canid specimens in this cluster may represent possible progenitors of the domestic dog. Given the position of the S805 haplotype, it may potentially represent a direct link from the putative progenitor (including Canis c.f. variabilis) to the domestic dog and modern wolf lineages.[43]

Ancestral dog

In 1999, a study emphasized that while molecular genetic data seem to support the origin of dogs from wolves, dogs may have descended from a now extinct species of canid whose closest living relative was the wolf.[25]

Goyet dog – 36,000 BP

Genus Canis, species indeterminate

In 2009, a study looked at 117 skulls of recent and fossil large canids. None of the 10 canid skulls from the Belgian caves of Goyet, Trou du Frontel, Trou de Nutons, and Trou de Chaleux could be classified, so the team took as their basic assumption that all of these canid samples were wolves.[52] The DNA sequence of 7 of the skulls indicated 7 unique haplotypes that represented ancient wolf lineages lost until now. The osteometric analysis of the skulls showed that one large canid fossil from Goyet (RBINS 2860) was clearly different from recent wolves, resembling most closely the Eliseevichi-1 dogs (15,000 years BP) and so was identified as a Paleolithic dog (see below).[50][53]

In November 2013, a DNA study sequenced 3 haplotypes from the ancient Belgium canids (the Goyet dog - Belgium 36,000 years BP cataloged as Canis species KF661079, and with Belgium 30,000 years BP KF661080 and 26,000 years BP KF661078 cataloged as Canis lupus) and found they formed the most diverging group. Although the cranial morphology of the Goyet dog has been interpreted as dog-like, it's mitochondrial DNA relation to other canids places it as an ancient sister-group to all modern dogs and wolves rather than a direct ancestor. Belgium 26,000 has been found to be uniquely large but was found not to be related to the Beringian wolf. This Belgium canid clade may represent a phenotypically distinct and not previously recognized population of gray wolf or the Goyet dog may represent an aborted domestication episode.[3]

Altai dog - 33,000 BP

Genus Canis, species indeterminate
33,000 year old skull of a dog-like canid found in the Altai Mountains. It has no direct descendants today

In 2011, a study looked at the well-preserved 33,000-year old skull and left mandible of a dog-like canid that was excavated from Razboinichya Cave in the Altai Mountains of southern Siberia (Central Asia). The morphology was compared to the skulls and mandibles of large Pleistocene wolves from Predmosti, Czech Republic dated 31,000 BP, modern wolves from Europe and North America, and prehistoric Greenland dogs from the Thule period (1,000 BP or later) to represent a large-sized but unimproved fully domestic dogs. "The Razboinichya Cave cranium is virtually identical in size and shape to prehistoric Greenland dogs" and not the ancient nor modern wolves. However, the lower carnassial tooth fell within the lower range of values for prehistoric wolves and was only slightly smaller than modern European wolves, and the upper carnassial tooth fell within the range of modern wolves. "We conclude, therefore, that this specimen may represent a dog in the very early stages of domestication, i.e. an incipient dog, rather than an aberrant wolf... The Razboinichya Cave specimen appears to be an incipient dog...and probably represents wolf domestication disrupted by the climatic and cultural changes associated with the Last Glacial Maximum".[54]

In March 2013, a DNA study sequenced the Altai dog with the data deposited in GenBank with accession number JX173682 and classified as Canis lupus familiaris (dog). The sample was compared with those of 72 extant dogs and 30 wolves (17 Old World and 13 New World), 35 prehistoric New World canids (including 2 from the Beringian wolf), and 4 coyotes. The sample was also compared to 3 ancient wolf teeth also found in the same cave (32,500 BP, 48,000 BP and 50,000 BP). "The analyses revealed that the unique haplotype of the Altai dog is more closely related to modern dogs and prehistoric New World canids than it is to contemporary wolves... This preliminary analysis affirms the conclusion that the Altai specimen is likely an ancient dog with shallow divergence from ancient wolves. These results suggest a more ancient history of the dog outside of the Middle East or East Asia." The haplotype groups closest to the Altai dog included such diverse breeds as the Tibetan Mastiff, Newfoundland, Chinese Crested, Cocker Spaniel and Siberian Husky. The study also stated "We stress the point that these analyses were limited to a single, maternally inherited locus and more sequence data would be needed to obtain a statistically well-supported phylogeny and unambiguously resolve the genetic relationship of the Altai specimen."[55]

In November 2013, a study looked at 18 fossil canids and compared these with the complete mitochondrial genome sequences from 49 modern wolves and 77 modern dogs. A more comprehensive analysis of the complete mtDNA found that the phylogenetic position of the Altai dog as being either dog or wolf was inconclusive and cataloged its sequence as Canis species (KF661092). The sequence strongly suggests a position at the root of a clade uniting two ancient wolf genomes, two modern wolves, as well as two dogs of Scandinavian origin, however the study does not support its recent common ancestry with the great majority of modern dogs. The study suggests that it may represent an aborted domestication episode.[3]

Paleolithic dog - 27,000 BP

Detailed DNA analysis yet to be conducted

In 2002, a study looked at 2 fossil skulls of large canids (references: MAE RAS 447 and ZIN RAS 23781) dated at 13,905 years BP that had been found buried within metres of what was once a mammoth-bone hut at the Upper Paleolithic site of Eliseevichi-1 in the Brayansk region of central Russia, and using an accepted morphologically-based definition of domestication declared them to be "Ice Age dogs".[56] The most complete dog skull was found in a hearth deposit, near a concentration of mammoth skulls. Its braincase has been perforated on the left and right side. Cut marks are present on the zygomatic and frontal bones. With exception of the canines and some premolars, all its teeth are missing. In addition the left and right carnassials were apparently removed by damaging the alveoli.[50] In 2013, a study recalibrated the age of the Eliseevichi-1 specimens to 15,000 BP with the genetic analysis of one skull deposited in GenBank with accession number KF661082 and classified as Canis lupus familiaris (dog).[3] It is the oldest known specimen recognized as belonging to a dog.

In 2009, a study proposed that there was a low frequency of recognized dog skulls in Upper Paleolithic sites because existing specimens had not yet been recognized as dogs. The study looked at these 2 prehistoric dog skulls in comparison to much earlier Late Pleistocene but morphologically similar fossil skulls that had been found across Europe, and proposed the much earlier specimens were Paleolithic dogs that were morphologically and genetically distinct from the Pleistocene wolves living in Europe at that time. The study looked at 117 skulls of recent and fossil large canids. Several skulls of fossil large canids from sites in Belgium, the Ukraine and Russia were examined using multivariate techniques to look for possible evidence of the presence of Paleolithic dogs that were separate from Pleistocene wolves. Reference groups constituted of the Eliseevichi-1 prehistoric dogs, and recent wolves and dogs. The osteometric analysis of the skulls indicated that the Paleolithic dogs fell outside the skull ranges of the Pleistocene wolf group and the modern wolf group, and were closer related to those of the Eliseevichi-1 prehistoric dog group. The fossil large canid from Goyet, Belgium (RBINS 2860), dated at 31,700 BP was clearly different from the recent wolves, resembling most closely the Eliseevichi 1 prehistoric dogs. The two Epigravettian Mezin, Ukraine (PM NASU 5490) and Mezhirich, Ukraine (PM NASU 4493) skulls are also identified as being Paleolithic dogs. Collagen analysis indicated that the Paleolithic dogs associated with human camp-sites (Eliseevichi 1, Mezin and Mezhirich) had been specifically eating reindeer, while other predator species in those locations and times had eaten a range of prey.[50]

Further studies have looked at wolf-like fossils from Paleolithic hunter-gatherer sites across Europe and propose to have identified Paleolithic dogs at Predmosti (Czech Republic 26,000-27,000 years BP), Kosteki-8 (Russia 23,000-27,700 years BP), Kosteki-1 (Russia 22,000-24,000 BP), Kosteki-17 (Russia Upper Paleolithic) and Verholenskaya (Russia late glacial).[57] In the human burial zone at the Predmosti site, 3 Paleolithic skulls were found that resemble those of a Siberian husky but they were larger and heavier than the modern husky. For one skull, "a large bone fragment is present between the upper and lower incisors that extends several centimetres into the mouth cavity. The size, thickness and shape of the fragment suggest that it could be a fragment of a bone of a large mammal, probably from a mammoth. The position of the bone fragment in the mouth and the articulated state of the lower jaw with the skull indicate that this mammoth bone fragment was inserted artificially into the mouth of the dog post-mortem." The morphology of some wolf-like fossils was such that they could not be assigned to either the Pleistocene wolf nor Paleolithic dog groups.[58]

In comparison to the Pleistocene wolf, the Paleolithic dog had a short skull length, short snout, a wide palate and braincase, relatively short and massive jaws, and a shorter carnassial length but these were larger than the modern dog. The mandible of the Paleolithic dog is more massive compared to the elongated mandible of the wolves, and presents a high frequency of crowded premolars and of a hook-like extension of the caudal border of the coronoid process. The Paleolithic dog had a mean body mass of 36–37 kg compared to Pleistocene wolves 42–44 kg and recent European wolves 41–42 kg.[50]

It has been proposed that based on the genetic evidence of the timeline and European location, the archaeological evidence of the Paleolithic dog remains being found at known European hunting camp-sites, and based on morphology and collagen analysis that their diet had been restricted, the Paleolithic dog was domesticated. It has been further proposed that the Paleolithic dog may have provided the stock from which early dogs came from.[50] There has been an ongoing debate in the scientific press about what the fossil remains of the Paleolithic "dog" might be, with some commenters declaring them as either wolves or a unique form of wolf. These include a first article proposing the Paleolithic dog,[50] its refutation,[59] a counter to the refutation,[60] a second article,[58] its refutation,[61] a third article that includes a counter to the refutation,[57] its refutation,[62] a counter to the refutation,[63] and another refutation.[64]

As the ancestor of the dog has not been positively identified by scientists, this debate continues.

See also Paleolithic dog

Second divergence

Gray wolf admixture

There was admixture between the ancestral dog, the ancestral modern gray wolf, and the golden jackal
The ancestral dog triverged into the Dingo, Basenji and Boxer lineages, and the ancestral modern gray wolf split into today's gray wolves
Dog breeds like this Tamaskan Dog look like wolves due to admixture.

In January 2014, a study analysed the whole-genome sequences of 3 wolves (Canis lupus) to represent the regions of Eurasia where domestication has been hypothesized to have taken place – Croatia (Europe), Israel (Middle-East), and China (East/South-East Asia), plus an Australian dingo and a Basenji, being divergent lineages to the reference boxer genome (that is available from GenBank with accession number AAEX01000000) and so maximize the odds to capture distinct alleles present in the earliest dogs. These lineages are also geographically distinct, with modern Basenjis tracing their ancestry to hunting dogs of western Africa, while dingoes are free-living semi-feral dogs of Australia that arrived there at least 3,500 years ago. The natural range of wolves has never extended this far south and due to geographic isolation, they are less likely to have overlapped and admixed with wolves in the recent past. For some analyses, data were leveraged from a companion study of 12 additional dog breeds.[2]

The data provided significant evidence of admixture between the Israeli wolf and the Basenji, the Israeli wolf and the boxer, and between the Chinese wolf and dingo. The Chinese wolf with dingo likely represents ancient admixture in Eastern Eurasia, and the Israeli wolf with Basenji and Boxer likely represents ancient admixture in Western Eurasia. The fact that these lineages have been geographically isolated from wolves in the recent past suggests that this gene flow was ancestral and has likely affected most dog lineages. There was significant gene flow between the golden jackal and the Israeli wolf, as well as the population ancestral to the dog and wolf samples.[2]

One test indicated that dogs and modern wolves form sister clades i.e. the dog is a sister to the modern wolf and they share a common ancestor. Supporting this, another test indicated that none of the sampled wolf populations is more closely related to dogs than any of the others, and dogs diverged from wolves at about the same time as wolves diverged from each other. This implies that the wolf population(s) from which dogs originated has gone extinct and the current wolf diversity from each region represents novel, younger wolf lineages.[2]

The data indicate that the golden jackal and the ancestor of the wolf/dog diverged 400,000 years ago. Dogs and wolves then diverged into the ancestral dog and the ancestral modern gray wolf. The ancestral modern gray wolf population triverged into the 3 populations studied. Not long after, the ancestral dog populations diverged into the dingo lineage, the basenji lineage and the reference boxer lineage. There was a 16-fold population bottleneck for dogs since this divergence.[2]

There was a 3-fold population decline for the 3 wolf samples since divergence, and appears to have occurred well in advance of direct extermination campaigns by humans and within the timeframe of environmental and biotic changes associated with the ending of the Pleistocene era i.e. changes in climate and prey, including megafaunal extinctions. This indicates that before the divergence of dogs from wolves that there was much more wolf diversity. The results support a recent divergence between dogs and wolves followed by a dramatic reduction in population size.[2]

The AMY2B (Alpha-Amylase 2B) is a gene that codes a protein that assists with the first step in the digestion of dietary starch and glycogen. An expansion of this gene in dogs would enable early dogs to exploit a starch-rich diet as they fed on refuse from agriculture. Data indicated that the wolves and dingo had just 2 copies of the gene, the Siberian Husky that is associated with hunter-gatherers had just 3–4 copies, whereas the Saluki that is associated with the Fertile Crescent where agriculture originated had 29 copies. The results show that on average, modern dogs have a high copy number of the gene whereas wolves and dingoes do not. The high copy number of AMY2B variants likely already existed as a standing variation in early domestic dogs, but expanded more recently with the development of large agriculturally based civilizations. This suggests that at the beginning of the domestication process, dogs may have been characterized by a more carnivorous diet than their modern-day counterparts, a diet held in common with early hunter-gatherers.[2]

Greenland sled dog carries 3.5% shared genetic material with the 35,000 years BP Taymyr wolf specimen.

Taimyr wolf admixture

There was admixture between Taimyr-1 and those breeds associated with high latitudes

In May 2015, a study compared the ancestry of the Taimyr-1 wolf lineage to that of dogs and gray wolves.

Comparison to the gray wolf lineage indicated that Taimyr-1 was basil to gray wolves from the Middle East, China, Europe and North America but sharing a substantial amount of history with the present-day gray wolves after their divergence from the coyote. This implies that the ancestry of the majority of gray wolf populations today stems from an ancestral population that lived less than 35,000 years ago but before the inundation of the Bering Land Bridge with the subsequent isolation of Eurasian and North American wolves.[1]:21

A comparison of the ancestry of the Taimyr-1 lineage to the dog lineage indicated that some modern dog breeds have a closer association with either the Gray wolf or Taimyr-1 due to admixture. The Sarloos wolfdog showed more association with the Gray wolf and this is in agreement with the documented historical crossbreeding with gray wolves in this breed. Taimyr-1 shared more alleles (i.e. gene expressions) with those breeds that are associated with high latitudes - the Siberian husky and Greenland dog that are also associated with arctic human populations, and to a lesser extent the Shar Pei and Finnish spitz. An admixture graph of the Greenland dog indicates a best-fit of 3.5% shared material, however an ancestry proportion ranging between 1.4% and 27.3% is consistent with the data. This indicates admixture between the Taimyr-1 population and the ancestral dog population of these 4 high-latitude breeds. These results can be explained either by a very early presence of dogs in northern Eurasia or by the genetic legacy of Taimyr-1 being preserved in northern wolf populations until the arrival of dogs at high latitudes. This introgression could have provided early dogs living in high latitudes with phenotypic variation beneficial for adaption to a new and challenging environment. It also indicates the ancestry of present-day dog breeds descends from more than one region.[1]:3-4

An attempt to explore admixture between Taimyr-1 and gray wolves produced unreliable results.[1]:23

Archaeological evidence

Archaeology locates the earliest cofirmed dog and human remains together 15,000 years ago at the Eliseevich-I site, Russian Plain, Russia.

Years BP Location Finding
400,000 Eurasia Lower Paleolithic sites including Boxgrove near Kent, England (400,000 years BP), Zhoukoudian in North China (300,000 BP), and Lazeret in southern France (150,000 years BP) have yielded wolf bones in close association with hominid bones. “The sites of occupation and hunting activities of humans and wolves must often have overlapped.”[32] We do not know if the co-location was the result of coincidence or a relationship.
200,000 Zhoukoudian cave system, China Small, extinct wolf skulls – Canis lupus variabilis. The skull differs from the typical wolf in much smaller size with a more slender muzzle and noticeably reduced or absent sagittal crest. In addition, the lower border of some Canis lupus variabilis mandibles is "strongly convex as in the dog".[51]:15 More recent researchers have proposed that Canis lupus variabilis may be an ancestor of the dog lineage.[65][66]:7 At the site, the small wolf's remains were in close proximity to Homo erectus pekinensis or Peking man. In 2015, a mitochondrial DNA analysis was conducted on 14 ancient canid remains from Arctic Siberia and suggests a genetic contribution from regional sources of wolves, including possibly Canis cf. variabilis, to the modern dog lineage. This was the first study to extract DNA material from Canis variabilis and it was thought to be widespread in Eurasia until 300,000 years ago.[67]
26,000 Chauvet cave, Vallon-Pont-d'Arc, Ardèche region, France 50-metre trail of footprints made by a boy of about ten years of age alongside those of a large canid. The size and position of the canid's shortened middle toe in relation to its pads indicates a dog rather than a wolf. The footprints have been dated by soot deposited from the torch the child was carrying. The cave is famous for its cave paintings.[68]
15,000 Eliseevich-I site, Russian Plain, Russia Two fossil dog skulls (references: MAE RAS 447 and ZIN RAS 23781). In 2002, a study looked at the fossil skulls of 2 large canids that had been found buried within metres of what was once a mammoth-bone hut at the Upper Paleolithic site of Eliseevichi 1 in the Brayansk region of central Russia, and using an accepted morphologically-based definition of domestication declared them to be "Ice Age dogs".[56] The Eliseevich-I skulls are very similar in shape to the Goyet skull (36,000 BP) and to the Epigravettian Mezin 5490 and Mezhirich dog skulls (13,500 BP). The remains of at least eight mammoth bone complexes and large quantities of worked ivory were discovered at this site.[50] Genetic analysis of one skull was deposited in GenBank with accession number KF661082 and classified as Canis lupus familiaris (dog).[3] See Paleolithic dog.
14,700 Bonn-Oberkassel, Germany Ancient dog mandible. Directly associated with a human double-grave of a 50-year-old man and a 20-25-year-old woman.[69] Genetic analysis deposited in GenBank with accession number KF661093 and classified as Canis lupus familiaris (dog).[3]
13,500 approx Mezin, Ukraine Ancient dog skull (PM NASU 5490) as well as ancient wolf specimens found at the site. Dated to the Epigravettian period (17,000–10,000 BP). The Mezin skull is very similar in shape to the Goyet skull (36,000 BP), the Eliseevich I dog skulls (15,000) and Mezhirich dog skull (13,500 BP). The Epigravettian Mezin site is well known for its round mammoth bone dwelling.[50]
13,500 approx Mezhirich, Ukraine Ancient dog skull (PM NASU 4493). Dated to the Epigravettian period (17,000–10,000 BP). The Mezhirich skull is very similar in shape to the Goyet skull (36,000 BP), the Eliseevich I dog skulls (15,000) and Mezin dog skull (13,500 BP). The Epigravettian Mazhirich site has four mammoth bone dwellings present.[50]
12,500 Karstein cave, Germany Ancient dog skull. Genetic analysis deposited in GenBank with accession number KF661094 and classified as Canis lupus familiaris (dog).[3]
12,000 Yakutia, Siberia Mummified carcass. The "Black Dog of Tumat" was found frozen into the ice core of an oxbow lake steep ravine at the middle course of the Syalaah River in Ust-Yana region. DNA analysis confirmed it as an early dog.[70]
12,000 Ain Mallaha (Eynan) and Hayonim terrace, Israel Three canid finds. A diminutive carnassial and a mandible, and a wolf or dog puppy skeleton buried with a human during the Natufian culture.[71]
9,200 Texas, USA Dog bone fragment. Found in Hinds Cave in southwest Texas. DNA analysis confirms the bone was from a dog whose ancestry was rooted in Eurasia.[72]
7,800 Jiahu site, China Eleven dog internments. Jaihu is a Neolithic site 22 kilometers north of Wuyang in Henan Province.[73]
7,425 Baikal region, Siberia, Russia Dog buried in a human burial ground. Additionally, a human skull was found buried between the legs of a "tundra wolf" dated 8,320 BP (but it does not match any known wolf DNA stored in Genbank). The evidence indicates that as soon as formal cemeteries developed in Baikal, some canids began to receive mortuary treatments that closely paralleled those of humans. One dog was found buried with four red deer canine pendants around its neck dated 5,770 BP. Many burials of dogs continued in this region with the latest finding at 3,760 BP, and they were all buried laying on their right side, and faced towards the east as did their humans. Some were buried with artifacts, e.g., stone blades, birch bark and antler bone.[74]
5,250 Skateholm, Sweden Cemeteries contained dogs among humans. A dog burial with an antler headdress and three flint blades was recovered at one of the sites.[75]


See also differences from wolves


The theory of dog domestication is based on a comparison between the dog and extant (i.e. living today) gray wolves, however one study highlighted a number of inconsistencies with this comparison and proposed that the ancestor of the dog appears more likely to have been a generalist canid and not the specialized gray wolf.[66] Another study proposed that the ancestor of Canis familiaris was a wild "Canis familiaris".[76] A genetic study recently indicated that the ancestor of the dog was not the extant gray wolf and that the fossil remains of the "wolf-like canid" ancestor have yet to be found.[3] Another found the same thing and that the dog, the Gray wolf and the now-extinct Taymyr wolf all diverged from a common ancestor 40,000 years ago i.e. without human intervention.[1] Readers should be aware of these inconsistencies with the theory before progressing further into this topic.


Domestication is a process by which a population of animals becomes adapted to man and the captive environment by some combination of genetic changes occurring over generations.[77] Domestication is an evolutionary process in which one population of a species is reproductively isolated from another intentionally by humans.[78] This reproductive isolation leads to an divergent adaptation and results in a specialization process. As a result of the changes in the selection pressures on the given species, the process of domestication produces evolutionary changes in certain aspects of the characteristic behavior of the domesticated species just as it affects the anatomy and morphology of the certain species as well. For example, such behavioral change in dogs is the decreased level of aggression, which is manifested in morphological changes such as teeth size.[79]

Self domestication

The first of the two main hypotheses of dog domestication is self-domestication by wolves. Some wolves moved into a symbiotic relationship with prehistoric humans. They scavenged on the remains of the prey animals left by the prehistoric people at the human settlements or the kill sites. Those wolves that were less anxious and aggressive thrived, continued to follow the prehistoric humans and colonized the human-dominated environments, generation after generation. Gradually, the first primitive dogs emerged from this group.[80][81][82]

Natural selection against aggression without human intervention

Dogs can infer the name of an object and have been shown to learn the names of over 1,000 objects. Dogs can follow the human pointing gesture; even nine week old puppies can follow a basic human pointing gesture without being taught. New Guinea Singing dogs, a half-wild proto-dog endemic to the remote alpine regions of New Guinea, as well as Dingoes in the remote outback of Australia are also capable of this. These examples demonstrate an ability to read human gestures that arose early in domestication and did not require human selection. "Humans did not develop dogs, we only fine-tuned them down the road."[83]:92 Similar to the chimpanzee, Bonobos are a close genetic cousin to humans. Unlike the chimpanzee, bonobos are not aggressive and do not participate in lethal intergroup aggression or kill within their own group. The most distinctive features of a bonobo are its cranium, which is 15% smaller than a chimpanzee's, and its less aggressive and more playful behavior. Dogs mirror these differences relative to wild wolves: a dog's cranium is 15% smaller than an equally heavy wolf's, and the dog is less aggressive and more playful. The guinea pig's cranium is 13% smaller than its wild cousin the cavie and domestic fowl show a similar reduction to their wild cousins. Possession of a smaller cranium for holding a smaller brain is a telltale sign of domestication. Bonobos appear to have domesticated themselves.[83]:104 In the "farm fox" experiment, humans selectively bred foxes against aggression which caused a domestication syndrome. The foxes were not selectively bred for smaller craniums and teeth, floppy ears, or skills at using human gestures but these traits were demonstrated in the friendly foxes. Natural selection favours those that are the most successful at reproducing, not the most aggressive. Selection against aggression made possible the ability to cooperate and communicate among foxes, dogs and bonobos. Perhaps it did the same thing for humans.[83]:114[84]


A genetic study has found a dog-wolf divergence time of greater than 15,000 years ago. An evolutionary scenario consistent with these results is that dog domestication was initiated close to the Last Glacial Maximum when hunter-gathers preyed on megafauna. Conceivably, proto-dogs might have taken advantage of carcasses left on site by early hunters, assisted in the capture of prey, or provided defense from large competing predators at kills.[3]

A leading evolutionary biologist has stated:

But if domestication occurred in association with hunter-gatherers, one can imagine wolves first taking advantage of the carcasses that humans left behind – a natural role for any large carnivore – and then over time moving more closely into the human niche through a co-evolutionary process. The idea of wolves following hunter-gatherers also helps to explain the eventual genetic divergence that led to the appearance of dogs. Wolves following the migratory patterns of these early human groups would have given up their territoriality and would have been less likely to reproduce with resident territorial wolves. We have an analog of this process today, in the only migratory population of wolves known existing in the tundra and boreal forest of North America. This population follows the barren-ground caribou during their thousand-kilometer migration. When these wolves return from the tundra to the boreal forest during the winter, they do not reproduce with resident wolves there that never migrate. We feel this is a model for domestication and the reproductive divergence of the earliest dogs from wild wolves. We know also that there were distinct wolf populations existing ten of thousands of years ago. One such wolf, which we call the megafaunal wolf, preyed on large game such as horses, bison and perhaps very young mammoths. Isotope data show that they ate these species, and the dog may have been derived from a wolf similar to these ancient wolves in the late Pleistocene of Europe....A dog from Belgium dates back approximately 36,000 years, and a group of dogs from Western Russia is approximately 15,000 years old.[85]
See also Megafaunal wolf


Some researchers argue that those wolves that were more successful at interacting with humans would pass these traits on to their offspring, eventually creating wolves with a greater propensity to be domesticated. The behavioral characteristic called "flight distance" represents how close an animal will allow humans (or anything else it perceives as dangerous) to get before it runs away. Animals with shorter flight distances will linger and feed when humans are close by. This behavioral trait would have been passed on to successive generations, and amplified, creating animals that are increasingly more comfortable around humans. To be able to eat in the presence of human beings is something that wild wolves can't do.[86] The "most social and least fearful" wolves were the ones who were kept around the human living areas, helping to breed those traits that are still recognized in dogs today.[87][88]

Against this proposition, wolves have been scavenging around human living areas, noted in Israel and Italy for centuries, without demonstrating any move towards domestication.[89]

Polychrome cave painting of a wolf-like canid painted 17,000 years ago, Font-de-Gaume, France.

Human domestication

The second of the two main hypotheses of dog domestication is domestication by humans. Paleolithic people actively selected wolf pups for several reasons: they could be used as pets, they could be kept for utilitarian, ceremonial and symbolic uses, as social storage, or combat and/or as living tools.[82][90][91][92][93] The most docile or interesting animals could have been permitted to reproduce.[91][92][32] After several generations of unconscious and later of conscious selection of human-defined behavioral traits, the first dogs emerged.[94]

Orphaned wolf-pups

Studies have shown that some wolf pups taken at an early age and reared by humans are easily tamed and socialized,[26]:140 and one study has demonstrated that adult wolves can be socialized.[26]:141 Some researchers propose that humans adopted orphaned wolf pups and breastfed them alongside human babies.[87][95] In Alaska and other northern areas where people still live close to wolves, wolf pups are sometimes captured and some become acceptable as pets or sled dogs, and once these breeding over generations would become more dog-like.[26]:pages55-56

Against this proposition, at the time of domestication near the LGM humans were already the top predator and had no need for wolves that would grow to eat 5 kilograms of meat per wolf per day at a time when food was very scarce. Starvation would have been a real threat to many carnivores in the Ice Age and competition for food would have been fierce.[83]:29 Other researchers attempting to socialize wolf pups after they reach 21 days of age found it very time-consuming and seldom practical or reliable in achieving success.[96]

Human selection

See also: Dog breeding

The "farm fox" experiment attempted to reenact how domestication may have occurred.[97] Researchers, working with farmed silver foxes selectively bred over 35 generations and 40 years for "tameability". The "domestic elite" foxes were tamer to humans than others but they also showed new physical traits even though the physical traits were not originally selected for. These include spotted or black-and-white coats, floppy ears, tails that curl over their backs, the barking vocalization and earlier sexual maturity. One researcher found that the migration of certain melanocytes (which determine colour) was delayed, resulting in a black and white 'star' pattern.

One criticism of this experiment was later made by the author, who stated that the living conditions of the foxes in the farm would have been very different to those of wolf puppies in Paleolithic camps.[98] A further criticism based on information obtained after the experiment's publication is that the definition of "tame" was changed at least once during the experiment, and that some of the foxes that were classified as neither tame nor aggressive also exhibited these changes, indicating that some factor other than human selection for tameness may have been at work during domestication.[89] When humans restrict dog's breeding diversity, another variable also comes into play that may have contributed to the change – inbreeding.[87]:30

In 2014, a whole genome study of the DNA differences between wolves and dogs found that dogs did not show a reduced fear response, they showed greater synaptic plasticity. Synaptic plasticity is widely believed to be the cellular correlates of learning and memory and this change may have altered the learning and memory abilities of dogs.[99]

Convergent evolution

Behavioral evidence

Convergent evolution is when distantly related species independently evolve similar solutions to the same problem. For example, fish, penguins and dolphins have each separately evolved flippers as solution to the problem of moving through the water. What has been found is something less frequently demonstrated: psychological convergence. Dogs have independently evolved to be cognitively more similar to humans than we are to our closest genetic relatives.[83]:60 Dogs have evolved specialized skills for reading human social and communicative behavior. These skills seem more flexible – and possibly more human-like – than those of other animals more closely related to humans phylogentically, such as chimpanzees, bonobos and other great apes. This raises the possibility that convergent evolution has occurred: both Canis familiaris and Homo sapiens might have evolved some similar (although obviously not identical) social-communicative skills – in both cases adapted for certain kinds of social and communicative interactions with human beings.[100]

The pointing gesture is a human-specific signal, is referential in its nature, and is a foundational building-block of human communication. Human infants acquire it weeks before the first spoken word.[101] In 2009, a study compared the responses to a range of pointing gestures by dogs and human infants. The study showed little difference in the performance of 2-year-old children and dogs, while 3-year-old children’s performance was higher. The results also showed that all subjects were able to generalize from their previous experience to respond to relatively novel pointing gestures. These findings suggest that dogs demonstrate a similar level of performance as 2-year-old children that can be explained as a joint outcome of their evolutionary history as well as their socialization in a human environment.[102]

Later studies support coevolution in that dogs can discriminate the emotional expressions of human faces,[103] most people can tell from a bark whether a dog was alone or being approached by a stranger, playing or being aggressive,[104] and able tell from a growl how big the dog is.[105]

Biological evidence

In 2013, a DNA sequencing study indicated that parallel evolution in humans and dogs is most apparent in the genes for digestion and metabolism, neurological process and cancer, likely as a result of shared selection pressures.[106][107]

In 2014, a study compared the hemoglobin levels of village dogs and people on the Chinese lowlands with those on the Tibetan plateau. It found the hemoglobin levels higher for both people and dogs in Tibet, suggesting that Tibetan dogs might share similar adaptive strategies as the Tibetan people. A population genetic analysis then showed a significant convergence between humans and dogs in Tibet.[108]

In 2015, a study found that when dogs and their owners interact, extended eye contact (mutual gaze) increases oxytocin levels in both the dog and its owner. As oxytocin is known for its role in maternal bonding, it is considered likely that this effect has supported the coevolution of human-dog bonding.[109]

The dog could have arisen only from animals predisposed to human society by lack of fear, attentiveness, curiosity, necessity, and recognition of advantage gained through collaboration....were not biological Audio-Animatronics born with a preprogrammed response...It is fair, I think, to say that the humans and wolves involved in the conversion were sentient, observant beings constantly making decisions about how they lived and what they did, based on the perceived ability to obtain at a given time and place what the needed to survive and thrive. They were social animals willing, even eager, to join forces with another animal to merge their sense of group with the others' sense and create an expanded super-group that was beneficial to both in multiple ways. They were individual animals and people involved, from our perspective, in a biological and cultural process that involved linking not only their lives but the evolutionary fate of their heirs n ways, we must assume, they could never have imagined. Does this thesis project too much self-awareness into the past? I doubt it. Powerful emotions were in play that many observers today refer as love – boundless, unquestioning love.

Lupification of humans

Isn’t it strange that, our being such an intelligent primate, we didn’t domesticate chimpanzees as companions instead? Why did we choose wolves even though they are strong enough to maim or kill us?[110]

Bison surrounded by gray wolf pack

In 2002, a study argued that immediate human ancestors and wolves may have domesticated each other through a strategic alliance that would change both respectively into humans and dogs. The effects of human psychology, hunting practices, territoriality and social behavior would have been profound.[111]

Marking of territory with signs such as pecked cupules, hand stencils and prints, abraded grooves, and finger impressions in once-soft mud are enduring signs used to mark occupation. They also became the first symbolic objects i.e. art. Wolves mark their territory with urine, however humans do not have the keen sense of smell as wolves and would have needed to use something more easily recognizable and enduring to mark their territory. Humans learned to mark their territory after watching wolves and dogs.[111]

Hunting large animals in packs is a distinctive wolf behavioral trait. There is no evidence of big game hunting in pre-sapiens groups, but big-game hunting is very typical of homo sapiens that, in addition to climate change, may have contributed to the extinction of many large mammals. Early humans moved from scavenging and small-game hunting to big-game hunting by living in larger, socially more-complex groups, learning to hunt in packs, and developing powers of cooperation and negotiation in complex situations. As these are characteristics of wolves, dogs and humans, it can be argued that these behaviors were enhanced once wolves and humans began to cohabit. Communal hunting lead to communal defense. Wolves actively patrol and defend their scent-marked territory, and perhaps humans had their sense of territoriality enhanced by living with wolves.[111]

New forms of bonding might assist in living in large, complex and varied social groups. One of the keys to recent human survival has been the negotiation of situations by forming partnerships. Strong bonds exist between same-sex wolves, dogs and humans – bonds less-fickle than exists between other same-sex animal pairs. Today, the most widespread form of inter-species bonding occurs between humans and dogs. The concept of friendship has ancient origins, but it may have been enhanced through the inter-species relationship to give a survival advantage.[111]

In 2003, a study compared the behavior and ethics of chimpanzees, wolves and humans. The sociality of humans' closest relative, the chimpanzee, appears as a frightful caricature of human egoism. Even in their maternal behavior, warmth and affection are reduced to nursing and the occasional comforting hug. Cooperation among group members is limited to occasional hunting episodes or the persecution of a competitor, always aimed for one's own advantage. The closest approximation to human morality they can find in nature is that of the gray wolf, Canis lupus. Wolves ability to cooperate in well-coordinated drives to hunt prey, carry items too heavy for an individual, provisioning not only their own young but also the other pack members, babysitting etc. are rivalled only by that of human societies. Similar forms of cooperation are observed in two closely related canids, the African Cape hunting dog and the Asian dhole, therefore it is reasonable to assume that canid sociality and cooperation are old traits that in terms of evolution predate human sociality and cooperation. Today's wolves may even be less social than their ancestors, as they have lost access to big herds of ungulates and now tend more toward a lifestyle similar to coyotes, jackals, and even foxes.[110]

There are a number of behavioral adaptions that make communal life possible:

Reindeer moved in large herds across the Mammoth steppe and were preyed upon by carnivores
  1. Social pack-forming canids are essentially monogamous. One pair breeds, but all members share food and parental care generously.
  2. They are aware of where the other pack members are and what they are doing, especially when running as a single group. When canids hunt in a pack, close attention and close cooperation allow them to operate as an integrated system.
  3. In wolves, each pack member can accept greater risks when attacking, because if injured the needy will be fed by the other pack members. Cooperation and risk sharing is not only among close relatives or mated pairs, but also lasting friendships among the same gender, which is a central feature of canid pack living.
  4. "Wolfing down" prey as quickly as possible by the pack ensures that other predators get no advantage. They later regurgitate large chunks for the pups and their baby-sitters.
  5. They constantly watch each other – they know who is who, who is where, and what they are doing.
  6. They deal with hierarchy with minimal bloodshed, with dominance and submission treated as a low-key affair.[110]

The Mammoth steppe was the Eurasian tundra and grass steppe ecosystem which once stretched from Spain to the far east of Siberia, and at times continued into North America. On this steppe the wolves ability to hunt in packs, to share risk fairly among pack members, and to cooperate moved them to the top of the food pyramid above lions, hyenas and bears. Some, but not all, wolves followed the great reindeer herds, eliminating the unfit, the weaklings, the sick and the aged and therefore improved the herd. These wolves had become the first pastoralists hundreds of thousands of years before humans also took to this role. The wolves advantage over their competitors was that they were able to keep pace with the herds, move fast and enduringly, and make the most efficient use of their kill by their ability to "wolf down" a large part of their quarry before other predators had detected the kill. The authors of the study propose that during the last ice age, some of our ancestors teamed up with those pastoralist wolves. Many of our ancestors remained gatherers and scavengers, or specialized as fish-hunters, hunter-gatherers, and the hunter-gardeners. However, some ancestors adopted the pastoralist wolves' lifestyle as herd followers and herders of reindeer, horses, and other hoofed animals. They harvested the best stock for themselves while the wolves kept the herd strong. These pastoralists later became herders and the dog.[110]

From a biologist's vantage point, the interwining process of hominization and canization makes sense only if viewed in terms of coevolution.[110]

Further reading

  • Derr, Mark. How the Dog Became the Dog: From Wolves to Our Best Friends (Penguin Group; 2011); book extensively cites research papers to support its propositions.
  • Hare, Brian & Woods, Venessa. The Genius of Dogs (2013 Penguin Publishing Group) Part 1, Chapters 2-5, pages 17–121; these several chapters focus on the cognitive evolution of the dog.
  • Morey, Darcy. Dogs: Domestication and the Development of a Social Bond (Cambridge University Press; 2010) 384 pages; uses zooarchaeology to explores ties between humans and canines over the past 15,000 years with a focus on the New World and Arctic regions.


  1. ^ a b c d e f g h i Skoglund, P.; Ersmark, E.; Palkopoulou, E.; Dalén, L. (2015). "Ancient Wolf Genome Reveals an Early Divergence of Domestic Dog Ancestors and Admixture into High-Latitude Breeds". Current Biology. doi:10.1016/j.cub.2015.04.019.  edit
  2. ^ a b c d e f g h Freedman, A. H.; Gronau, I.; Schweizer, R. M.; Ortega-Del Vecchyo, D.; Han, E.; Silva, P. M.; Galaverni, M.; Fan, Z.; Marx, P.; Lorente-Galdos, B.; Beale, H.; Ramirez, O.; Hormozdiari, F.; Alkan, C.; Vilà, C.; Squire, K.; Geffen, E.; Kusak, J.; Boyko, A. R.; Parker, H. G.; Lee, C.; Tadigotla, V.; Siepel, A.; Bustamante, C. D.; Harkins, T. T.; Nelson, S. F.; Ostrander, E. A.; Marques-Bonet, T.; Wayne, R. K.; Novembre, J. (2014). "Genome Sequencing Highlights the Dynamic Early History of Dogs". PLoS Genetics 10 (1): e1004016. doi:10.1371/journal.pgen.1004016. PMC 3894170. PMID 24453982. 
  3. ^ a b c d e f g h i j k l m Thalmann, O.; Shapiro, B.; Cui, P.; Schuenemann, V. J.; Sawyer, S. K.; Greenfield, D. L.; Germonpré, M. B.; Sablin, M. V.; López-Giráldez, F.; Domingo-Roura, X.; Napierala, H.; Uerpmann, H-P.; Loponte, D. M.; Acosta, A. A.; Giemsch, L.; Schmitz, R. W.; Worthington, B.; Buikstra, J. E.; Druzhkova, A.; Graphodatsky, A. S.; Ovodov, N. D.; Wahlberg, N.; Freedman, A. H.; Schweizer, R. M.; Koepfli, K.-.P.; Leonard, J. A.; Meyer, M.; Krause, J.; Pääbo, S.; Green, R. E.; Wayne, R. K. (2013). "Complete Mitochondrial Genomes of Ancient Canids Suggest a European Origin of Domestic Dogs". Science 342 (6160): 871–74. doi:10.1126/science.1243650. 
  4. ^ a b Intergovernmental Panel on Climate Change (UN). "IPCC Fourth Assessment Report: Climate Change 2007 - Palaeoclimatic Perspective". The Nobel Foundation. 
  5. ^ doi:10.1126/science.1172873
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  6. ^ doi:10.1126/science.1172873
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  7. ^ doi:10.1038/382060a0
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  8. ^ Jonathan Adams. "Europe during the last 150,000 years". Oak Ridge National Laboratory, Oak Ridge, USA. 
  9. ^ doi:10.1007/978-1-4757-5202-1_11
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  10. ^ Dale Guthrie, R. (2004). "Radiocarbon evidence of mid-Holocene mammoths stranded on an Alaskan Bering Sea island". Nature 429 (6993): 746–749. doi:10.1038/nature02612. PMID 15201907. 
  11. ^ doi:10.1671/0272-4634(2003)23[260:LPSOTS]2.0.CO;2
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  12. ^ Barnett, R.; Shapiro, B.; Barnes, I. A. N.; Ho, S. Y. W.; Burger, J.; Yamaguchi, N.; Higham, T. F. G.; Wheeler, H. T.; Rosendahl, W.; Sher, A. V.; Sotnikova, M.; Kuznetsova, T.; Baryshnikov, G. F.; Martin, L. D.; Harington, C. R.; Burns, J. A.; Cooper, A. (2009). "Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity". Molecular Ecology 18 (8): 1668–1677. doi:10.1111/j.1365-294X.2009.04134.x. PMID 19302360.  edit
  13. ^ doi:10.1016/j.quascirev.2014.12.020
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  14. ^ doi:10.1126/science.aac4315
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  15. ^ White, T. D.; Asfaw, B.; Degusta, D.; Gilbert, H.; Richards, G. D.; Suwa, G.; Clark Howell, F. (2003). "Pleistocene Homo sapiens from Middle Awash, Ethiopia". Nature 423 (6941): 742–7. doi:10.1038/nature01669. PMID 12802332. 
  16. ^ "A Human Journey:Migration Routes". The genographic project. National Geographic Society. 2015. Retrieved 2015. 
  17. ^ Benazzi, S.; Douka, K.; Fornai, C.; Bauer, C. C.; Kullmer, O.; Svoboda, J. Í.; Pap, I.; Mallegni, F.; Bayle, P.; Coquerelle, M.; Condemi, S.; Ronchitelli, A.; Harvati, K.; Weber, G. W. (2011). "Early dispersal of modern humans in Europe and implications for Neanderthal behaviour". Nature 479 (7374): 525. doi:10.1038/nature10617. 
  18. ^ Higham, T.; Compton, T.; Stringer, C.; Jacobi, R.; Shapiro, B.; Trinkaus, E.; Chandler, B.; Gröning, F.; Collins, C.; Hillson, S.; o’Higgins, P.; Fitzgerald, C.; Fagan, M. (2011). "The earliest evidence for anatomically modern humans in northwestern Europe". Nature 479 (7374): 521. doi:10.1038/nature10484. 
  19. ^ doi:10.1038/35092552
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  20. ^ "Mamontovaya Kurya:an enigmatic, nearly 40000 years old Paleolithic site in the Russian Arctic" (PDF). 
  21. ^ doi:10.1126/science.1085219
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  22. ^ Tamm, E.; Kivisild, T.; Reidla, M.; Metspalu, M.; Smith, D. G.; Mulligan, C. J.; Bravi, C. M.; Rickards, O.; Martinez-Labarga, C.; Khusnutdinova, E. K.; Fedorova, S. A.; Golubenko, M. V.; Stepanov, V. A.; Gubina, M. A.; Zhadanov, S. I.; Ossipova, L. P.; Damba, L.; Voevoda, M. I.; Dipierri, J. E.; Villems, R.; Malhi, R. S. (2007). Carter, Dee, ed. "Beringian Standstill and Spread of Native American Founders". PLoS ONE 2 (9): e829. doi:10.1371/journal.pone.0000829. PMC 1952074. PMID 17786201. 
  23. ^ Balter, M (4 July 2013). "Farming Was So Nice, It Was Invented at Least Twice". Science. 
  24. ^ Darwin, Charles (1868). "1". The Variation of Animals and Plants under Domestication 1. John Murray, London. 
  25. ^ a b c doi:10.1002/(sici)1521-1878(199903)21:3<247::aid-bies9>;2-z
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  26. ^ a b c d Scott, John Paul (1965). Genetics and the Social Behavior of the Dog:The Classic Study. University of Chicago Press. pp. 55–56. ISBN 978-0-226-74338-7. 
  27. ^ Boyko, Adam; Boykob, Ryan H.; Boykob, Corin M.; Parkerc, Heidi G.; Castelhanod, Marta; Corey, L.; Degenhardt, J. D.; Auton, A.; Hedimbi, M.; Kityo, R.; Ostrander, E. A.; Schoenebeck, J.; Todhunter, R. J.; Jones, P.; Bustamante, C. D. (2009-08-18). "Complex population structure in African village dogs and its implications for inferring dog domestication history". PNAS 106 (33): 13903–13908. doi:10.1073/pnas.0902129106. PMID 19666600. 
  28. ^ Pang et al. (September 1, 2009). "mtDNA Data Indicate a Single Origin for Dogs South of Yangtze River, Less Than 16,300 Years Ago, from Numerous Wolves". Molecular Biology and Evolution. doi:10.1093/molbev/msp195. 
  29. ^ Wayne, Robert K. (1993). "Molecular evolution of the dog family". Trends in Genetics 9 (6): 218–224. doi:10.1016/0168-9525(93)90122-X. PMID 8337763. 
  30. ^ Wozencraft, W. Christopher (1993). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Animal Species of the World:A Taxonomic and Geographic Reference 1 (2 ed.). Washington, D.C.: Smithsonian Institution Press. p. 576. ISBN 9780801882210. 
  31. ^ Gentry A, Clutton-Brock J, Groves C P (2004). "The naming of wild animal species and their domestic derivatives" 31 (5). Journal of Archaeological Science. pp. 645–651. doi:10.1016/j.jas.2003.10.006. 
  32. ^ a b c Clutton-Brock, J., 1995. Origins of the dog: domestication and early history. In:Serpell, J. (Ed.), The Domestic Dog, its Evolution, Behaviour and Interactions with People. Cambridge University Press, Cambridge, pp. 7e20.
  33. ^ Carles Vilà, Peter Savolainen, Jesús E. Maldonado, Isabel R. Amorim, John E. Rice, Rodney L. Honeycutt, Keith A. Crandall, Joakim Lundeberg, Robert K. Wayne (1997). "Multiple and Ancient Origins of the Domestic Dog" 276 (5319doi=10.1126/science.276.5319.1687). Science. pp. 1687–1689. 
  34. ^ a b Pilot, M. G.; Branicki, W.; Jędrzejewski, W. O.; Goszczyński, J.; Jędrzejewska, B. A.; Dykyy, I.; Shkvyrya, M.; Tsingarska, E. (2010). "Phylogeographic history of grey wolves in Europe". BMC Evolutionary Biology 10: 104. doi:10.1186/1471-2148-10-104. PMC 2873414. PMID 20409299. 
  35. ^ Tamm, E.; Kivisild, T.; Reidla, M.; Metspalu, M.; Smith, D. G.; Mulligan, C. J.; Bravi, C. M.; Rickards, O.; Martinez-Labarga, C.; Khusnutdinova, E. K.; Fedorova, S. A.; Golubenko, M. V.; Stepanov, V. A.; Gubina, M. A.; Zhadanov, S. I.; Ossipova, L. P.; Damba, L.; Voevoda, M. I.; Dipierri, J. E.; Villems, R.; Malhi, R. S. (2007). Carter, Dee, ed. "Beringian Standstill and Spread of Native American Founders". PLoS ONE 2 (9): e829. doi:10.1371/journal.pone.0000829. PMC 1952074. PMID 17786201. 
  36. ^ Leonard, J. A.; Vilà, C; Fox-Dobbs, K; Koch, P. L.; Wayne, R. K.; Van Valkenburgh, B (2007). "Megafaunal extinctions and the disappearance of a specialized wolf ecomorph" (PDF). Current Biology 17 (13): 1146–50. doi:10.1016/j.cub.2007.05.072. PMID 17583509. 
  37. ^ Germonpré, M.; Sablin, M. V.; Stevens, R. E.; Hedges, R. E. M.; Hofreiter, M.; Stiller, M.; Després, V. R. (2009). "Fossil dogs and wolves from Palaeolithic sites in Belgium, the Ukraine and Russia: Osteometry, ancient DNA and stable isotopes". Journal of Archaeological Science 36 (2): 473. doi:10.1016/j.jas.2008.09.033.  edit
  38. ^ Hofreiter, M.; Barnes, I. (2010). "Diversity lost: Are all Holarctic large mammal species just relict populations?". BMC Biology 8: 46. doi:10.1186/1741-7007-8-46. PMC 2858106. PMID 20409351. 
  39. ^ a b doi:10.1016/j.cub.2007.06.031
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  40. ^ doi:10.1016/j.jas.2012.06.016
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  41. ^ PMID 15643947 (PubMed)
    Citation will be completed automatically in a few minutes. Jump the queue or expand by hand
  42. ^ "Ancient wolf genome pushes back dawn of the dog". Nature. 
  43. ^ a b c doi:10.1371/journal.pone.0125759
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  44. ^ Vilà C; Savolainen P; Maldonado JE; Amorim IR; Rice JE; Honeycutt RL; Crandall KA; Lundeberg J; Wayne RK (1997). "Multiple and ancient origins of the domestic dog". Science 276 (5319): 1687–1689. doi:10.1126/science.276.5319.1687. PMID 9180076. 
  45. ^ Savolainen, Peter; Zhang, Ya-ping; Luo, Jing; Lundeberg, Joakim; Leitner, Thomas (2002-11-22). "Genetic Evidence for an East Asian Origin of Domestic Dogs". Science 298 (5598): 1610–1613. Bibcode:2002Sci...298.1610S. doi:10.1126/science.1073906. PMID 12446907. 
  46. ^ Boyko, Adam; Boykob, Ryan H.; Boykob, Corin M.; Parkerc, Heidi G.; Castelhanod, Marta; Corey, L.; Degenhardt, J. D.; Auton, A.; Hedimbi, M.; Kityo, R.; Ostrander, E. A.; Schoenebeck, J.; Todhunter, R. J.; Jones, P.; Bustamante, C. D. (2009-08-18). "Complex population structure in African village dogs and its implications for inferring dog domestication history". PNAS 106 (33): 13903–13908. Bibcode:2009PNAS..10613903B. doi:10.1073/pnas.0902129106. PMC 2728993. PMID 19666600. 
  47. ^ Pang et al. (September 1, 2009). "mtDNA Data Indicate a Single Origin for Dogs South of Yangtze River, Less Than 16,300 Years Ago, from Numerous Wolves". Molecular Biology and Evolution. Retrieved 2010-01-07. 
  48. ^ vonHoldt, Bridgett; Lohmueller, Kirk E.; Han, Eunjung; Parker, Heidi G.; Quignon, Pascale; Degenhardt, Jeremiah D.; Boyko, Adam R.; Earl, Dent A.; Auton, Adam; Reynolds, Andy; Bryc, Kasia; Brisbin, Abra; Knowles, James C.; Mosher, Dana S.; Spady, Tyrone C.; Elkahloun, Abdel; Geffen, Eli; Pilot, Malgorzata; Jedrzejewski, Wlodzimierz; Greco, Claudia; Randi, Ettore; Bannasch, Danika; Wilton, Alan; Shearman, Jeremy; Musiani, Marco; Cargill, Michelle; Jones, Paul G.; Qian, Zuwei et al. (2010-03-17). "Genome-wide SNP and haplotype analyses reveal a rich history underlying dog domestication". Nature 464 (7290): 898–902. Bibcode:2010Natur.464..898V. doi:10.1038/nature08837. PMC 3494089. PMID 20237475. 
  49. ^ Ding, Z-L et al. (2011-11-23). "Origins of domestic dog in Southern East Asia is supported by analysis of Y-chromosome DNA". Heredity 108 (5): 507–14. doi:10.1038/hdy.2011.114. PMC 3330686. PMID 22108628Advance online publication 
  50. ^ a b c d e f g h i j Germonpré, Mietje; Sablin, Mikhail V.; Stevens, Rhiannon E.; Hedges, Robert E.M.; Hofreiter, Michael; Stiller, Mathias; Despre´s, Viviane R. (2009). "Fossil dogs and wolves from Palaeolithic sites in Belgium, the Ukraine and Russia: osteometry, ancient DNA and stable isotopes". Journal of Archaeological Science 36 (2): 473–490. doi:10.1016/j.jas.2008.09.033. 
  51. ^ a b Pei, W.C. (1934). The carnivora from locality 1 of Choukoutien. Palaeontologia Sinica, Series C, vol. 8, Fascicle 1. Geological Survey of China, Beijing. pp. 1–45. 
  52. ^ Pat Shipman (2011). The Animal Connection: A New Perspective on What Makes Us Human. W W Norton & Co New York. p. 218. 
  53. ^ "Goyet skull photo - Royal Belgium Institute of Natural Sciences". 
  54. ^ Ovodov, Nikolai D.; Crockford, Susan J.; Kuzmin, Yaroslav V.; Higham, Thomas F. G.; Hodgins, Gregory W. L.; van der Plicht, Johannes (July 28, 2011). "A 33,000-Year-Old Incipient Dog from the Altai Mountains of Siberia: Evidence of the Earliest Domestication Disrupted by the Last Glacial Maximum". PLoS ONE 6 (pages =): e22821. doi:10.1371/journal.pone.0022821. 
  55. ^ doi:10.1371/journal.pone.0057754
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  56. ^ a b Sablin, Mikhail V.; Khlopachev, Gennady A. (2002). "The Earliest Ice Age Dogs:Evidence from Eliseevichi I" (PDF). Wenner-Gren Foundation for Anthropological Research. Retrieved 10 January 2015. 
  57. ^ a b Germonpré, Mietje; Laznickova-Galetova, Martina; Losey, Robert J.; Raikkonen, Jannikke; Sablin, Mikhail V. (2014). "Large canids at the Gravettian Predmostí site, the Czech Republic:The mandible". Quaternary International xxx: 1–19. doi:10.1016/j.quaint.2014.07.012. 
  58. ^ a b Germonpré, Mietje; Laznickova-Galetova, Martina; Sablin, Mikhail V. (2012). "Palaeolithic dog skulls at the Gravettian Predmosti site, the Czech Republic". Journal of Archaeological Science 39 (1): 184–202. doi:10.1016/j.jas.2011.09.022. 
  59. ^ Crockford SJ, Kuzmin YV (2012) Comments on Germonpre et al., Journal of Archaeological Science 36, 2009 "Fossil dogs and wolves from Palaeolithic sites in Belgium, the Ukraine and Russia: osteometry, ancient DNA and stable isotopes"
  60. ^ Germonpré, Mietje; Sablin, MV; Despres, V; Hofreiter, M; Laznickova-Galetova, M et al. (2013). "Palaeolithic dogs and the early domestication of the wolf: a reply to the comments of Crockford and Kuzmin (2012)". Journal of Archaeological Science 40 (1): 786–792. doi:10.1016/j.jas.2012.06.016. 
  61. ^ Boudadi-Maligne, Myriam; Escarguel, Gilles (2014). "A biometric re-evaluation of recent claims for Early Upper Palaeolithic wolf domestication in Eurasia". Journal of Archaeological Science (Elsevier Ltd) 45: 80–89. doi:10.1016/j.jas.2014.02.006. 
  62. ^ Morley, Darcy F. (2014). "In search of Paleolithic dogs: a quest with mixed results". Journal of Archaeological Science 52: 300–307. doi:10.1016/j.jas.2014.08.015. 
  63. ^ Germonpré, Mietje; Sablin, Mikhail V.; Laznickova-Galetova, Martina; Despre´s, Viviane R.; Stevens, Rhiannon E.; Stiller, Mathias; Hofreiter, Michael (2015). "Palaeolithic dogs and Pleistocene wolves revisited: a reply to Morey (2014)". Journal of Archaeological Science 54: 210–216. doi:10.1016/j.jas.2014.11.035. 
  64. ^ Drake, Abby Grace; Coquerelle, Michael; Colombeau, Guillaume (5 February 2015). "3D morphometric analysis of fossil canid skulls contradicts the suggested domestication of dogs during the late Paleolithic". Scientific Reports 5 (2899): 8299. doi:10.1038/srep08299. 
  65. ^ Lawrence, B. (1967). "Early Domestic Dogs". Zeitschrift für Säugetierkunde 32: 44–59. 
  66. ^ a b Koler-Matznick, Janice (2002). "The Origin of the Dog Revisited". Anthrozoos 15 (2): 98–118. doi:10.2752/089279302786992595. 
  67. ^ doi: 10.1371/journal.pone.0125759
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  68. ^ Garcia, M.A. (2005). "Ichnologie ge´ne´ rale de la grotte Chauvet". Bulletin de la Socie´te´ pre´historique française 102 (1): 103–108. doi:10.3406/bspf.2005.13341. 
  69. ^ Morey, Darcy, ed. (2010). Dogs: Domestication and the Development of a Social Bond [1]. Cambridge, UK: Cambridge University Press. p. 24. ISBN 9780521757430. 
  70. ^ Fedorova, Anna (August 18, 2014). "Yakut scientists found remains of ancient dog". Yakutia Today. Retrieved February 19, 2015.  see also [2]
  71. ^ SJM Davis, FR Valla (7 December 1978). "Evidence for domestication of the dog 12,000 years ago in the Natufian of Palestine". Nature 276 (5688): 608–610. doi:10.1038/276608a0. 
  72. ^ Raul Y. Tito, Samuel L. Belknap, Kristin D. Sobolik, Robert C. Ingraham, Lauren M. Cleeland, Cecil M. Lewis Jr. (August 2011). "Brief communication: DNA from early Holocene American dog". American Journal of Physical Anthropology 145 (4): 653–657. doi:10.1002/ajpa.21526. 
  73. ^ Susan J. Crockford, A Practical Guide to In Situ Dog Remains for the Field Archaeologist, 2009 [3]
  74. ^ Losey, RJ; Bazaliiskii, VI; Garvie-Lok, S; Germonpré, M; Leonard, JA; Allen, AL; Anne Katzenberg, M; Sablin, MV (2011). "Canids as persons: Early Neolithic dog and wolf burials, Cis-Baikal, Siberia". Journal of Anthropological Archaeology 30 (2): 174–189. doi:10.1016/j.jaa.2011.01.001. 
  75. ^ "The Materiality of Death Bodies, burials, beliefs" Edited by Fredrik Fahlander Terje Oestigaard, 2008 BAR International Series 17682008 [4]
  76. ^ Manwell, C. & C. M. A. Baker. 1983. Origin of the dog: from wolf or wild Canis familiaris? Speculations in Science and Technology 6 (3): 213–224.
  77. ^ Grandin, T. and M.J. Deesing (1998). Genetics and the Behavior of domestic animals. Academic Press, San Diego, CA. ISBN 978-0-12-394586-0. 
  78. ^ kretchmer, KR; Fox, MW (1 February 1975). "Effects of domestication on animal behaviour". Vet Rec 96 (5): 102–108. doi:10.1136/vr.96.5.102. PMID 1090069. 
  79. ^ lakatos, Gabriella (2011). "Evolutionary approach to communication between humans and dogs". Ann Ist Super Sanita 47 (4): 373–377. doi:10.4415/ANN_11_04_08. 
  80. ^ Crockford, Susan (2000). Crockford, Susan, ed. A commentary on dog evolution: regional variation, breed development and hybridization with wolves. Archaeopress BAR International Series 889. pp. 11–20. ISBN 978-1841710891. 
  81. ^ Coppinger, Raymond; Coppinger, Lorna (2001). Dogs: A Startling New Understanding of Canine Origin, Behavior & Evolution. Scribner New York. ISBN 0684855305. 
  82. ^ a b Russell, Nerissa (2012). Social Zooarchaeology: Humans and Animals in Prehistory. Cambridge University Press. ISBN 978-0-521-14311-0. 
  83. ^ a b c d e Hare, Brian & Woods, Venessa (2013). The Genius of Dogs. Penguin Publishing Group. 
  84. ^ PMID 16061417 (PubMed)
    Citation will be completed automatically in a few minutes. Jump the queue or expand by hand
  85. ^ Wolpert, Stuart (November 14, 2013), "Dogs likely originated in Europe more than 18,000 years ago, UCLA biologists report", UCLA News Room, retrieved December 10, 2014 
  86. ^ "Human Stars, The Animal Attraction". 
  87. ^ a b c Derr, Mark (2004). Dog's Best Friend:Annals of the Dog-Human Relationship. Chicago: University of Chicago Press. ISBN 0-226-14280-9. 
  88. ^ Dale, Steve (January 22, 2014). "Dogs are smarter than we think, and some are downright brilliant". Chicago Tribune. 
  89. ^ a b c Derr, Mark (2011). How the Dog Became the Dog: From Wolves to Our Best Friends. Penguin Group USA. ISBN 1-59020-700-9. 
  90. ^ Crabtree, P.J.; Campana, D.V. (1987). "A new model for the domestication of the dog". Mus. Appl. Sci. Cent. Archaeol. J 4: 98–102. 
  91. ^ a b Germonpre, M., 2010. Comments. In: Shipman, P. (Ed.), The Animal Connection and Human Evolution, CA Forum on Theory in Anthropology. Curr. Anthropol. 51, pp. 527e528.
  92. ^ a b Serpell, J., 1989. Pet-keeping and animal domestication: a reappraisal. In: Clutton-Brock (Ed.), The Walking Larder: Patterns of Domestication, Pastoralism and Predation. Unwin-Hyman, London, pp. 10e20.
  93. ^ Shipman, P (2010). "The animal connection and human evolution". Curr. Anthropol 51: 519–538. doi:10.1086/653816. 
  94. ^ Trut, L., Oskina, I., Kharlamova, A., 2009. Animal evolution during domestication:the domesticated fox as a model" Bioessays 31, 349e360.
  95. ^ Grandin, Temple; Johnson, Catherine (2005). Animals in Translation. New York, New York: Scribner. p. 87. ISBN 0-7432-4769-8. 
  96. ^ Klinghammer, Erich; Goodmann, Patricia Ann (1987). "Chapter 2: Socialization and management of wolves in captivity". In Frank, Harry. Man and Wolf: Advances, Issues, and Problems in Captive Wolf Research. Dr W. Junk Publishers. pp. 31–61 [36]. ISBN 90-6193-614-4. 
  97. ^ Trut, Lyudmila N. "Early Canid Domestication: The Farm-Fox Experiment: Foxes Bred for Tamability in a 40-year Experiment Exhibit Remarkable Transformation That Suggest an Interplay between Behavioral Genetics and Development." American Scientist 87.No. 2 (MARCH-APRIL 1999) (1999): 160–69. JSTOR. Web. 1 Jan. 2014. | url=
  98. ^ Trut, L.N., Plyusnina, I.Z., Oskina, I.N., 2004. An experiment on fox domestication and debatable issues of evolution of the dog. Russ. J. Genet. 40, 644e655.
  99. ^ doi:10.1093/gbe/evu245
    This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand
  100. ^ Hare B, Tomasello M. 2005. Human-like social skills in dogs? Trends in cognitive sciences. 9: 439-444 [5]
  101. ^ Butterworth, George (2003). "Pointing is the royal road to language for babies". 
  102. ^ Lakatos, Gabriella (2009). "A comparative approach to dogs’ (‘'Canis familiaris’’) and human infants’ comprehension of various forms of pointing gestures". Animal Cognition 12: 621–631. doi:10.1007/s10071-009-0221-4. 
  103. ^ Corsin A. Müller1, Kira Schmitt, Anjuli L.A. Barber, Ludwig Huber (2 July 2015). "Dogs can discriminate the emotional expressions of human faces". Current Biology 25 (5): 601–605. doi:10.1016/j.cub.2014.12.055. 
  104. ^ Brian Hare, Vanessa Woods (8 February 2013), "What Are Dogs Saying When They Bark? [Excerpt]", Scientific America, retrieved 17 March 2015 
  105. ^ Katherine Sanderson (23 May 2008), "Humans can judge a dog by its growl", Nature, doi:10.1038/news.2008.852, retrieved 17 March 2015  research available here
  106. ^ Wang, G. D.; Zhai, W.; Yang, H. C.; Fan, R. X.; Cao, X.; Zhong, L.; Wang, L.; Liu, F.; Wu, H.; Cheng, L. G.; Poyarkov, A. D.; Poyarkov Jr, N. A.; Tang, S. S.; Zhao, W. M.; Gao, Y.; Lv, X. M.; Irwin, D. M.; Savolainen, P.; Wu, C. I.; Zhang, Y. P. (2013). "The genomics of selection in dogs and the parallel evolution between dogs and humans". Nature Communications 4: 1860. doi:10.1038/ncomms2814. 
  107. ^ Cossins, Dan (May 16, 2013), "Dogs and Human Evolving Together", The Scientist, retrieved January 12, 2014 
  108. ^ Wang, G. -D.; Fan, R. -X.; Zhai, W.; Liu, F.; Wang, L.; Zhong, L.; Wu, H.; Yang, H. -C.; Wu, S. -F.; Zhu, C. -L.; Li, Y.; Gao, Y.; Ge, R. -L.; Wu, C. -I.; Zhang, Y. -P. (2014). "Genetic Convergence in the Adaptation of Dogs and Humans to the High-Altitude Environment of the Tibetan Plateau". Genome Biology and Evolution 6 (8): 2122. doi:10.1093/gbe/evu162. 
  109. ^ Miho Nagasawa, Shouhei Mitsui, Shiori En, Nobuyo Ohtani, Mitsuaki Ohta, Yasuo Sakuma, Tatsushi Onaka, Kazutaka Mogi, Takefumi Kikusui (17 April 2015). "Oxytocin-gaze positive loop and the coevolution of human-dog bonds". Science 348 (6232). pp. 333–336. doi:10.1126/science.1261022. 
  110. ^ a b c d e Schleidt, Wolfgang M.; Shalter, Michael D. (2003). "Co-evolution of Humans and Canids: An Alternative View of Dog Domestication: Homo Homini Lupus?" (PDF). Evolution and Cognition 9 (1): 57–72. 
  111. ^ a b c d Paul Taçon, Colin Pardoe (2002). "Dogs make us human". Nature Australia (Australian Museum) 27 (4): 52–61.  Journal no longer published