Origin of the domestic dog
The origin of the domestic dog includes the dog's evolutionary divergence from the wolf, its domestication, and its development into dog types and dog breeds. The dog is a member of the genus Canis, which forms part of the wolf-like canids, and was the first species and the only large carnivore to have been domesticated. The dog and the extant gray wolf are sister taxa, as modern wolves are not closely related to the population of wolves that was first domesticated.
The genetic divergence between dogs and wolves occurred between 40,000–20,000 years ago, just before or during the Last Glacial Maximum. This timespan represents the upper time-limit for the commencement of domestication because it is the time of divergence and not the time of domestication, which occurred later. The domestication of animals commenced over 15,000 years ago, beginning with the grey wolf (Canis lupus) by nomadic hunter-gatherers. The archaeological record and genetic analysis show the remains of the Bonn–Oberkassel dog buried beside humans 14,200 years ago to be the first undisputed dog, with disputed remains occurring 36,000 years ago. It was not until 11,000 years ago that people living in the Near East entered into relationships with wild populations of aurochs, boar, sheep, and goats.
Where the domestication of the dog took place remains debated, with the most plausible proposals spanning Western Europe, Central Asia and East Asia. This has been made more complicated by the recent proposal that an initial wolf population split into East and West Eurasian groups. These two groups, before going extinct, were domesticated independently into two distinct dog populations between 14,000 and 6,400 years ago. The Western Eurasian dog population was gradually and partially replaced by East Asian dogs introduced by humans at least 6,400 years ago. This proposal is also debated.
- 1 Canid and human evolution
- 2 Divergence from wolves
- 3 Dog domestication
- 3.1 Time of domestication
- 3.2 Socialization
- 3.3 Commensal pathway
- 3.4 Post-domestication gene flow
- 3.5 Positive selection
- 3.6 Natural selection
- 3.7 Dog and human convergent evolution
- 4 First dogs
- 5 References
- 6 Bibliography
- 7 Further reading
Canid and human evolution
Six million years ago, towards the close of the Miocene era, the earth's climate gradually cooled. This would lead to the glaciations of the Pliocene and the Pleistocene, which are commonly referred to as the Ice Age. In many areas, forests and savannahs were replaced with steppes or grasslands, and only those species of creature that adapted to these changes would survive.
In southern North America, small woodland foxes grew bigger and better adapted to running, and by the late Miocene the first of the genus Canis had arisen - the ancestors of coyotes, wolves and the domestic dog. In eastern Africa, a split occurred among the large primates. Some remained in the trees, while others came down from the trees, learned to walk upright, developed larger brains, and in the more open country learned to avoid predators while becoming predators themselves. The ancestors of humans and dogs would ultimately meet in Eurasia.
Human hunter-gatherers did not live in fear of nature and knew that they posed a formidable risk to any potential predators. Today, the Ju'wasi people of Namibia share their land with prides of lions. Both species coexist with respect and without fear or hostility in a relationship that may go back to the dawn of modern humans. The lion is a much larger and far more dangerous predator than the wolf. Early modern humans entering Eurasia and first encountering packs of wolves may have been assisted in living among them because of the traditional beliefs of their African ancestors. In historical times, mutual respect and cooperation with canids can be found in the stories and traditions of the indigenous peoples of Siberia, East Asia, North America, and Australia.
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 in ways, we must assume, they could never have imagined.
Divergence from wolves
The domestication of animals commenced over 15,000 years before present (YBP), beginning with the grey wolf (Canis lupus) by nomadic hunter-gatherers. It was not until 11,000 YBP that people living in the Near East entered into relationships with wild populations of aurochs, boar, sheep, and goats. A domestication process then began to develop. The grey wolf most likely followed the commensal pathway to domestication. When, where, and how many times wolves may have been domesticated remains debated because only a small number of ancient specimens have been found, and both archaeology and genetics continue to provide conflicting evidence. The most widely accepted, earliest dog remains date back 15,000 YBP to the Bonn–Oberkassel dog. Earlier remains dating back to 30,000 YBP have been described as Paleolithic dogs, however their status as dogs or wolves remains debated. Recent studies indicate that a genetic divergence occurred between dogs and wolves 20,000-40,000 YBP, however this is the upper time-limit for domestication because it represents the time of divergence and not the time of domestication.
Time of genetic divergence
The date estimated for the evolutionary divergence of a domestic lineage from a wild one does not necessarily indicate the start of the domestication process but it does provide an upper boundary. The divergence of the lineage that led to the domestic horse from the lineage that led to the modern Przewalski's horse is estimated to have occurred around 45,000 YBP but the archaeological record indicates 5,500 YBP. The variance can be due to modern wild populations not being the direct ancestor of the domestic ones, or to a divergence caused by changes in the climate, topography, or other environmental influences. The evolutionary divergence time for the wolf and dog is indicated to have occurred somewhere between 20,000-60,000 YBP but this does not imply that domestication occurred during this period.
During the Late Pleistocene glaciation, a vast mammoth steppe stretched from Spain eastwards across Eurasia and over the Bering land bridge into Alaska and the Yukon. The Late Pleistocene was characterized by a series of severe and rapid climate oscillations with regional temperature changes of up to 16 °C (61 °F), which has been correlated with megafaunal extinctions. There is no evidence of megafaunal extinctions at the height of the Last Glacial Maximum, indicating that increasing cold and glaciation were not factors. Multiple events appear to have caused the rapid replacement of one species by another one within the same genus, or one population by another within the same species, across a broad area. As some species became extinct, so too did the predators that depended on them.
The origin of dogs is couched in the paleobiogeography of wolf populations during the Late Pleistocene. The earliest fossils of Canis lupus were found in what was once eastern Beringia at Old Crow, Yukon, Canada and at Cripple Creek Sump, Fairbanks, Alaska. The age is not agreed but could date 1 million YBP. Considerable morphological diversity existed among grey wolves by the Late Pleistocene. These are regarded as having been more cranio-dentally robust than modern grey wolves, often with a shortened rostrum, the pronounced development of the temporalis muscle, and robust premolars. It is proposed that these features were specialized adaptations for the processing of carcass and bone associated with the hunting and scavenging of Pleistocene megafauna. Compared with modern wolves, some Pleistocene wolves showed an increase in tooth breakage that is similar to that seen in the extinct dire wolf. This suggests that these either often processed carcasses, or that they competed with other carnivores and needed to quickly consume their prey. The frequency and location of tooth fractures found in these wolves compared with the modern spotted hyena indicates that these wolves were habitual bone crackers.
|Grey wolf divergence and timing|
|Phylogenetic tree - extant grey wolf populations, with divergence times calculated using an assumed mutation rate of Skoglund (0.4x10−8) or [Lindblad-Toh] (1.0x10−8). The dog is a genetically divergent subspecies of the grey wolf, with substantial divergence between the dog lineages as nearly as distinct from one another as wolves are from dogs, which may reflect more admixture and wolf ancestry retained in their genome.|
Genetic studies indicate that the gray wolf is the closest living relative of the dog, with no evidence of any other canine species having contributed. Attempting to reconstruct the dog's lineage through the phylogenetic analysis of DNA sequences from modern dogs and wolves has given conflicting results for several reasons. Firstly, studies indicate that an extinct Late Pleistocene wolf is the nearest common ancestor to the dog, with modern wolves not being the dog's direct ancestor. Secondly, the genetic divergence between the dog and modern wolves occurred over a short period of time, so that the time of the divergence is difficult to date (referred to as incomplete lineage sorting). This is complicated further by the cross-breeding that has occurred between dogs and wolves since domestication (referred to as post-domestication gene flow). Finally, there have been only tens of thousands of generations of dogs since domestication, so that the number of mutations between the dog and the wolf are few and this makes the timing of domestication difficult to date.
In 2013, the whole genome sequencing of modern dogs and wolves indicated a divergence time of 32,000 YBP. In 2014, another study indicated 16,000-11,000 YBP. The first draft genome sequence of a Pleistocene canid was published in 2015. This Taymyr Peninsula wolf belonged to a population that had diverged from the ancestors of both modern wolves and dogs. Radiocarbon dating indicates its age to be 35,000 YBP, and this age could then be used to re-calibrate the wolf's mutation rate, indicating that the genetic divergence between dogs and wolves occurred before the Last Glacial Maximum, between 40,000–27,000 YBP. When this mutation rate was applied to the timing of the 2014 study, that study gave the same result of 40,000–27,000 YBP.
Place of genetic divergence
Most genetic studies conducted over the last two decades were based on modern dog breeds and extant wolf populations, with their findings dependent on a number of assumptions. These studies assumed that the extant wolf was the ancestor of the dog, did not consider genetic admixture between wolves and dogs, nor the impact of incomplete lineage sorting. These pre-genomic studies have suggested an origin of dogs in Southeast Asia, East Asia, Central Asia, the Middle East, or Europe. More recently, the field of Paleogenomics applies the latest molecular technologies to fossil remains that still contain useful DNA.
In 2013, a study sequenced the complete mitochondrial genomes of 18 fossil canids and these were then compared with the mitochondrial genomes from modern wolves and dogs in a phylogenetic tree. The study found that there exists more genetic variation between the 18 fossil canids than exists between all modern wolves and dogs. The analyses included the oldest fossils proposed as early dogs dating back as far as 36,000 YBP. However, these Goyet Cave canids were found to have formed an ancient and extinct sister group to modern dogs and wolves. The authors propose that these dog-like fossils were either an early domestication attempt that left no descendants among modern dogs, or that they were a specialized wolf ecomorph whose morphological features were part of the Late Pleistocene wolf diversity.
An earlier study of the Razboinichya Cave canid that was based on a segment of its mitochondrial DNA (mDNA) had concluded that it was an early dog. This fossil was included for the analysis of its complete mitochondrial genome and on the phylogenetic tree was assigned a much more basal position. The study inferred from the tree that dogs and wolves split 32,000-19,000 YBP and therefore the beginning of domestication occurred in the time of hunter-gatherers rather than in the time of farmers. The tree also confirmed that the ancient dogs of the Americas originated in Eurasia.
The phylogenetic analyses revealed that three of the four major mDNA clades of dogs relate most closely to the ancient canids from Europe rather than those from China or the Middle East, which supports a European origin of modern dogs. No modern wolf population related closer to dogs than the ancient canids from Europe, indicating that the wolf population that was the ancestor of the dog is extinct.
Arctic northeastern Siberia
In 2015, a study recovered mDNA from ancient canid specimens that were discovered in arctic northeastern Siberia (which was once western Beringia). These specimens included the mandible of a 360,000-400,000 YBP Canis c.f. variabilis (where c.f. is a Latin term meaning uncertain). Phylogenetic analyses of these canids revealed nine mDNA haplotypes not detected before. The Canis c.f. variabilis specimen clustered with other wolf samples from across Russia and Asia. The mDNA haplotypes of one 8,750 YBP specimen and some 28,000 YBP specimens matched with those of geographically widely-spread modern dogs. One 47,000 YBP canid was distinct from wolves but was only a few mutations away from those haplotypes found in modern dogs. The authors concluded that the structure of the modern dog gene pool was contributed to from ancient Siberian wolves and possibly from Canis c.f. variabilis.
Dogs show both ancient and modern lineages. The ancient lineages appear most in Asia but least in Europe because the Victorian era development of modern dog breeds used little of the ancient lineages. All dog populations (breed, village, and feral) show some evidence of genetic admixture between modern and ancient dogs. Some ancient dog populations that once occupied Europe and the New World no longer exist. This implies that some ancient dog populations were entirely replaced and others admixed over a long period of time. European dog populations have undergone extensive turnover during the last 15,000 years which has erased the genomic signature of early European dogs, the genetic heritage of the modern breeds has become blurred due to admixture, and there was the possibility of past domestication events that had gone extinct or had been largely replaced by more modern dog populations.
In 2016, a study compared the mitochondrial DNA and whole-genome sequences of a worldwide panel of modern dogs, the mDNA sequences of 59 ancient European dog specimens dated 14,000-3,000 YBP, and the nuclear genome sequence of a dog specimen that was found in the Late Neolithic passage grave at Newgrange, Ireland and radiocarbon dated at 4,800 YBP. A genetic analysis of the Newgrange dog showed that it was male, did not possess genetic variants associated with modern coat length nor color, was not as able to process starch as efficiently as modern dogs but more efficiently than wolves, and showed ancestry from a population of wolves that could not be found in other dogs nor wolves today. As the taxonomic classification of the "proto-dog" Paleolithic dogs as being either dogs or wolves remains controversial, they were excluded from the study. The phylogenetic tree generated from mDNA sequences found a deep division between the Sarloos wolfdog and all other dogs, indicating that breed's recent deriving from the German Shepherd and captive gray wolves. The next largest division was between eastern Asian dogs and western Eurasian (Europe and the Middle East) dogs that had occurred between 14,000-6,400 YBP, with the Newgrange dog clustering with the western Eurasian dogs.
The Newgrange and ancient European dog mDNA sequences could be largely assigned to mDNA haplogroups C and D but modern European dog sequences could be largely assigned to mDNA haplogroups A and B, indicating a turnover of dogs in the past from a place other than Europe. As this split dates older than the Newgrange dog this suggests that the replacement was only partial. The analysis showed that most modern European dogs had undergone a population bottleneck (reduction) which can be an indicator of travel. The archaeological record shows dog remains dating over 15,000 YBP in western Eurasia, over 12,500 YBP in eastern Eurasia, but none older than 8,000 YBP in Central Asia. The study proposes that dogs may have been domesticated separately in both eastern and western Eurasia from two genetically distinct and now extinct wolf populations. East Eurasian dogs then made their way with migrating people to western Europe between 14,000-6,400 YBP where they partially replaced the dogs of Europe. Two domestication events in western Eurasia and eastern Eurasia has recently been found for the domestic pig.
The hypothesis is that two genetically different, and possibly now extinct, wolf populations were domesticated independently in eastern and western Eurasia to produce paleolithic dogs. The eastern Eurasian dogs then dispersed westward alongside humans, reaching western Europe 6,400–14,000 years ago where they partially replaced the western paleolithic dogs. A single domestication is thought to be due to chance, however dual domestication on different sides of the world is unlikely to have happened randomly and it suggests that external factors - an environmental driver - may have forced wolves to work together with humans for survival. It is possible that wolves took advantage of resources that humans had, or humans may have been introduced to wolves in an area in which they didn't previously live.
The study indicates that the western Eurasian wolf and dog populations genetically diverged 20,000-60,000 YBP. Immediately after this divergence, the dog population outnumbered the wolf population, and later the dog population underwent a population reduction to be much lower.
Two origins disputed
In 2017, a study compared the nuclear genome sequences of three ancient dog specimens from Germany and Ireland with sequences from over 5,000 dogs and wolves. These Neolithic dog specimens included a dog sample from the Early Neolithic site in Herxheim, Germany dated 7,000 YBP, one from the Late Neolithic site of Kirschbaum (Cherry Tree) Cave near Forchheim, Germany dated 4,700 YBP, and a dog from Newgrange, Ireland dated 4,800 YBP. The study found that modern European dogs descended from their Neolithic ancestors with no evidence of a population turnover. There was evidence of a single dog-wolf divergence occurring between 36,900-41,500 YBP, followed by a divergence between Southeast Asian and Western Eurasian dogs 17,500-23,900 YBP and this indicates a single dog domestication event occurring between 20,000-40,000 YBP. The 3 dogs indicated ancestry that could be found in South East Asian dogs. Additionally, the Cherry Tree Cave dog showed ancestry that could be found in the Middle East, India and Central Asia. The study did not support a dual domestication event, and detected admixture between the ancestors of modern European and Southeast Asian dogs.
A 2018 study of mDNA sequences shows that the pre-Neolithic dogs of Europe all fell under haplogroup C. The Neolithic and Post-Neolithic dogs from Southeastern Europe that are associated with farmers fell under haplogroup D. In Western and Northern Europe, haplogroup D became diluted into the native dog population. This implies that haplogroup D arrived in Europe 9,000 years ago from the Near East along with pigs, cows, sheep, and goats. Later in 2018, another study looked at the y-chromosome male lineage of the ancient fossils of the Herxheim, Kirschbaum, and Newgrange dogs along with other canines. The study identified six major dog yDNA haplogroups, of which two of these include the majority of modern dogs. The Newgrange dog fell into the most commonly occurring of these haplogroups. The two ancient German dogs fell into a haplogroup commonly found among dogs from the Middle East and Asia, with the Kirschbaum dog sharing a common male lineage with the extant Indian wolf. The study concluded that at least 2 different male haplogroups existed in ancient Europe, and that the dog male lineage diverged from its nearest common ancestor shared with the gray wolf sometime between 68,000-151,000 YBP.
The questions of when and where dogs were first domesticated have taxed geneticists and archaeologists for decades. Identifying the earliest dogs is difficult because the key morphological characters that are used by zooarchaeologists to differentiate domestic dogs from their wild wolf ancestors (size and position of teeth, dental pathologies, and size and proportion of cranial and postcranial elements) were not yet fixed during the initial phases of the domestication process. The range of natural variation among these characters that may have existed in ancient wolf populations, and the time it took for these traits to appear in dogs, are unknown.
Early dog specimens
There are a number of recently discovered specimens which are proposed as being Paleolithic dogs, however their taxonomy is debated. These have been found in either Europe or Siberia and date 40,000-17,000 YBP. They include Hohle Fels in Germany, Goyet Caves in Belgium, Predmosti in the Czech Republic, and four sites in Russia: Razboinichya Cave, Kostyonki-8, Ulakhan Sular, and Eliseevichi 1. Paw-prints from Chauvet Cave in France dated 26,000 YBP are suggested as being those of a dog, however these have been challenged as being left by a wolf.
|40,000-35,000||Hohle Fels, Schelklingen, Germany||Paleolithic dog|
|36,500||Goyet Caves, Mozet, Belgium||Paleolithic dog|
|33,500||Razboinichya Cave, Altai Mountains, Central Asia||Paleolithic dog|
|33,500-26,500||Kostyonki-8, Voronezh, Russia||Paleolithic dog|
|31,000||Predmostí, Moravia, Czech Republic||Paleolithic dog|
|26,000||Chauvet Cave, Vallon-Pont-d'Arc, France||Paw-prints|
|17,200||Ulakhan Sular, northern Yakutia, Siberia||Paleolithic dog|
|17,000-16,000||Eliseevichi-I site, Bryansk Region, Russian Plain, Russia||Paleolithic dog|
There are also a number of later proposed Paleolithic dogs whose taxonomy has not been confirmed. These include a number of specimens from Germany (Kniegrotte, Oelknitz, Teufelsbrucke), Switzerland (Monruz, Kesslerloch, Champre-veyres-Hauterive), and Ukraine (Mezin, Mezhirich). A set of specimens dating 15,000-13,500 YBP have been confidently identified as domesticated dogs, based on their morphology and the archaeological sites in which they have been found. These include Spain (Erralla), France (Montespan, Le Morin, Le Closeau, Pont d’Ambon), and Germany (Bonn-Oberkassel). After this period, the remains of domesticated dogs have been identified from archaeological sites across Eurasia.
Possible dog domestication between 40,000-15,000 years ago is not clear due to the debate over what the Paleolithic dog specimens represent. This is due to the flexibility of genus Canis morphology, and the close morphological similarities between Canis lupus and Canis familiaris. It is also due to the scarcity of Pleistocene wolf specimens available for analyses and so their morphological variation is unknown. Habitat type, climate, and prey specialization greatly modify the morphological plasticity of grey wolf populations, resulting in a range of morphologically, genetically, and ecologically distinct wolf morphotypes. With no baseline to work from, zooarchaeologists find it difficult to be able to differentiate between the initial indicators of dog domestication and various types of Late Pleistocene wolf ecomorphs, which can lead to the mis-identification of both early dogs and wolves. Additionally, the ongoing prehistoric admixture with local wolf populations during the domestication process may have led to canids that were domesticated in their behavior but wolflike in their morphology. Attempting to identify early tamed wolves, wolfdogs, or proto-dogs through morphological analysis alone may be impossible without the inclusion of genetic analyses.
A domestication study looked at the reasons why the archeological record that is based on the dating of fossil remains often differed from the genetic record contained within the cells of living species. The study concluded that our inability to date domestication is because domestication is a continuum and there is no single point where we can say that a species was clearly domesticated using these two techniques. The study proposes that changes in morphology across time and how humans were interacting with the species in the past needs to be considered in addition to these two techniques.
..."wild" and "domesticated" exist as concepts along a continuum, and the boundary between them is often blurred — and, at least in the case of wolves, it was never clear to begin with.— Raymond Pierotti
|“||... Remove domestication from the human species, and there's probably a couple of million of us on the planet, max. Instead, what do we have? Seven billion people, climate change, travel, innovation and everything. Domestication has influenced the entire earth. And dogs were the first. For most of human history, we're not dissimilar to any other wild primate. We're manipulating our environments, but not on a scale bigger than, say, a herd of African elephants. And then, we go into partnership with this group of wolves. They altered our relationship with the natural world. ...||”|
|— Greger Larson|
The earlier association of dogs with humans may have allowed dogs to have a profound influence on the course of early human history and the development of civilization. However, the timing, geographic locations, and ecological conditions that led to dog domestication are not agreed.
There is clear evidence that dogs were derived from gray wolves during the initial phases of domestication and that no other canine species was involved. The wolf population(s) that were involved are likely to be extinct. Despite numerous genetic studies of both modern dogs and ancient dog remains, there is no firm consensus regarding either the timing or location(s) of domestication, the number of wolf populations that were involved, or the long-term effects domestication has had on the dog's genome.
Genetic studies suggest a domestication process commencing over 25,000 YBP, in one or several wolf populations in either Europe, the high Arctic, or eastern Asia. The remains of large carcasses left by human hunter-gatherers may have led some wolves into entering a migratory relationship with humans. This could have led to their divergence from those wolves that remained in the one territory. A closer relationship between these wolves — or proto-dogs — and humans may have then developed, such as hunting together and mutual defence from other carnivores and other humans. Around 10,000 YBP agriculture was developed resulting in a sedentary lifestyle, along with phenotype divergence of the dog from its wolf ancestors, including variance in size. In the Victorian era, directed human selection developed the modern dog breeds, which resulted in a vast range of phenotypes. Each of these domestication phases have left their mark on the dog's genome.
Time of domestication
An apex predator is a predator that sits on the top trophic level of the food chain, while a mesopredator sits further down the food chain and is dependent on smaller animals. Towards the end of the Pleistocene era, most of today's apex predators were mesopredators and this included the wolf. During the ecological upheaval associated with the close of the Late Pleistocene, one type of wolf population rose to become today's apex predator and another joined with humans to become an apex consumer.
In August 2015, a study undertook an analysis of the complete mitogenome sequences of 555 modern and ancient dogs. The sequences showed an increase in the population size approximately 23,500 YBP, which broadly coincides with the proposed separation of the ancestors of dogs and present-day wolves before the Last Glacial Maximum (refer first divergence). A ten-fold increase in the population size occurred after 15,000 YBP, which may be attributable to domestication events and is consistent with the demographic dependence of dogs on the human population.
Humans and wolves both exist in complex social groups. How humans and wolves got together remains unknown. One view holds that domestication as a process that is difficult to define. The term was developed by anthropologists with a human-centric view in which humans took wild animals (ungulates) and bred them to be "domestic", usually in order to provide improved food or materials for human consumption. That term may not be appropriate for a large carnivore such as the dog. This alternate view regards dogs as being either socialized and able to live among humans, or unsocialized. There exists today dogs that live with their human families but are unsocialized and will threaten strangers defensively and aggressively no different from a wild wolf. There also exists a number of cases where wild wolves have approached people in remote places, attempting to initiate play and to form companionship. One such notable wolf was Romeo, a gentle black wolf that formed relationships with the people and dogs of Juneau, Alaska. This view holds that before there could have been domestication of the wolf, there had to have been its socialization.
- See further: Convergent evolution between dogs and humans
The dog is a classic example of a domestic animal that likely traveled a commensal pathway into domestication. The dog was the first domesticant, and was domesticated and widely established across Eurasia before the end of the Pleistocene, well before cultivation or the domestication of other animals. It may have been inevitable that the first domesticated animal came from the order of carnivores as these are less afraid when approaching other species. Within the carnivores, the first domesticated animal would need to exist without an all-meat diet, possess a running and hunting ability to provide its own food, and be of a controllable size to coexist with humans, indicating the family Canidae, and the right temperament:p166 with wolves being among the most gregarious and cooperative animals on the planet.
- See further: Commensal pathway
Ancient DNA supports the hypothesis that dog domestication preceded the emergence of agriculture and was initiated close to the Last Glacial Maximum 27,000 YBP when hunter-gatherers preyed on megafauna, and when 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 kill-sites. Wolves were probably attracted to human campfires by the smell of meat being cooked and discarded refuse in the vicinity, first loosely attaching themselves and then considering these as part of their home territory where their warning growls would alert humans to the approach of outsiders. The wolves most likely drawn to human camps were the less-aggressive, subdominant pack members with lowered flight response, higher stress thresholds, less wary around humans, and therefore better candidates for domestication. The earliest sign of domestication in dogs was the neotenization of skull morphology and the shortening of snout length that results in tooth crowding, reduction in tooth size, and a reduction in the number of teeth, which has been attributed to the strong selection for reduced aggression. This process may have begun during the initial commensal stage of dog domestication, even before humans began to be active partners in the process.
A maternal mitochondrial, paternal Y chromosome, and microsatellite assessment of two wolf populations in North America and combined with satellite telemetry data revealed significant genetic and morphological differences between one population that migrated with and preyed upon caribou, and another territorial ecotype population that remained in a boreal coniferous forest. Though these two populations spend a period of the year in the same place, and though there was evidence of gene flow between them, the difference in prey–habitat specialization has been sufficient to maintain genetic and even coloration divergence. A study has identified the remains of a population of extinct Pleistocene Beringian wolves with unique mitochondrial signatures. The skull shape, tooth wear, and isotopic signatures suggested these were specialist megafauna hunters and scavengers that became extinct while less specialized wolf ecotypes survived. Analogous to the modern wolf ecotype that has evolved to track and prey upon caribou, a Pleistocene wolf population could have begun following mobile hunter-gatherers, thus slowly acquiring genetic and phenotypic differences that would have allowed them to more successfully adapt to the human habitat.
Even today, the wolves on Ellesmere Island do not fear humans, which is thought to be due to them seeing humans so little, and they will approach humans cautiously, curiously and closely.
- See further: Megafaunal wolf
Post-domestication gene flow
Since domestication, dogs have traveled alongside humans across most of the planet, often hybridizing with local wild canids. This has resulted in complex patterns of ancient and recent admixture among both the wild and the domestic canids.
Some studies have found greater diversity in the genetic markers of dogs from East and Central Asia compared to Europe and have concluded that dogs originated from these regions, despite no archaeological evidence to support the conclusions. One reason for these discrepancies is the sustained admixture between different dog and wolf populations across the Old and New Worlds over at least the last 10,000 years, which has blurred the genetic signatures and confounded efforts of researchers at pinpointing the origins of dogs. Another reason is that none of the modern wolf populations are related to the Pleistocene wolves that were first domesticated. In other words, the extinction of the wolves that were the direct ancestors of dogs has muddied efforts to pinpoint the time and place of dog domestication.
- See further: Post-domestication gene flow
|mDNA (maternal) ancestry of the Dog|
|Phylogenetic classification based on the entire mitochondrial genomes of the domestic dog. These resolve into 6 mDNA Haplogroups, most indicated as a result of male dog/female wolf hybridization.|
There is evidence of admixture between dog and regional wolf populations, except on the Tibetan Plateau and in the New World wolves. This admixture has occurred throughout history and as dogs expanded across the landscape. There are some dog populations that show recent admixture with wolves.
Phylogenetic analysis shows that modern dog mDNA haplotypes resolve into four monophyletic clades with strong statistical support, and these have been designated by researchers as clades A-D. Other studies that included a wider sample of specimens have reported two rare East Asian clades E-F with weaker statistical support. In 2009, a study found that haplogroups A, B and C included 98% of dogs and are found universally distributed across Eurasia, indicating that they were the result of a single domestication event, and that haplogroups D, E, and F were rare and appeared to be the result of regional hybridization with local wolves post-domestication. Haplogroups A and B contained subclades that appeared to be the result of hybridization with wolves post-domestication, because each haplotype within each of these subclades was the result of a female wolf/male dog pairing.
Haplogroup D: Derived from post-domestication wolf–dog hybridization in subclade d1 (Scandinavia) and d2 (South-West Asia). The northern Scandinavian subclade d1 hybrid haplotypes originated 480-3,000 YBP and are found in all Sami-related breeds: Finnish Lapphund, Swedish Lapphund, Lapponian Herder, Jamthund, Norwegian Elkhound and Hällefors Elkhound. The maternal wolf sequence that contributed to them has not been matched across Eurasia and its branch is phylogenetically rooted in the same sequence as the Altai dog (not a direct ancestor). The subclade d2 hybrid haplotypes are found in 2.6% of South-West Asian dogs.
Haplogroup F: Derived from post-domestication wolf–dog hybridization in Japan. A study of 600 dog specimens found only one dog whose sequence indicated hybridization with the extinct Japanese wolf.
It is not known whether this hybridization was the result of humans selecting for phenotypic traits from local wolf populations or the result of natural introgression as the dog expanded across Eurasia.
In 2018, a study found a small amount of dog ancestry in 62% of Eurasian wolf specimens looked at, that hybridization had occurred across a wide number of timescales and not just recently, however in contrast there was almost no admixture detected in the North American specimens. There was introgression of the male dog into the wolf, but also one hybrid detected which was the result of a male wolf crossed with a female dog. Wolves have maintained their phenotype differences from the dog, which indicates low-frequency hybridization. The conclusion is that phenotype is no indication of "purity" and the definition of pure wolves is ambiguous. Free-ranging dogs across Eurasia show introgression from wolves. Another study found that the β-defensin gene responsible for the black coat of North American wolves was the result of a single introgression from dogs in the Yukon dated between 1,600-7,200 years ago. The study proposes that early Native American dogs were the source.
Taimyr wolf admixture
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 basal to gray wolves from the Middle East, China, Europe and North America but shared 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.: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 Saarloos wolfdog showed more association with the gray wolf, which 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, although 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 four 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 that the ancestry of present-day dog breeds descends from more than one region.:3–4
An attempt to explore admixture between Taimyr-1 and gray wolves produced unreliable results.:23
As the Taimyr wolf had contributed to the genetic makeup of the Arctic breeds, a later study suggested that descendants of the Taimyr wolf survived until dogs were domesticated in Europe and arrived at high latitudes where they mixed with local wolves, and these both contributed to the modern Arctic breeds. Based on the most widely accepted oldest zooarchaeological dog remains, domestic dogs most likely arrived at high latitudes within the last 15,000 years. The mutation rates calibrated from both the Taimyr wolf and the Newgrange dog genomes suggest that modern wolf and dog populations diverged from a common ancestor between 20,000 and 60,000 YBP. This indicates that either dogs were domesticated much earlier than their first appearance in the archaeological record, or they arrived in the Arctic early, or both.
Charles Darwin recognized the small number of traits that made domestic species different from their wild ancestors. He was also the first to recognize the difference between conscious selective breeding in which humans directly select for desirable traits, and unconscious selection where traits evolve as a by-product of natural selection or from selection on other traits. Domestic animals have variations in coat color as well as texture, dwarf and giant varieties, and changes in their reproductive cycle, and many others have tooth crowding and floppy ears.
Although it is easy to assume that each of these traits was uniquely selected for by hunter-gatherers and early farmers, beginning in 1959 Dmitry Belyayev tested the reactions of silver foxes to a hand placed in their cage and selected the tamest, least aggressive individuals to breed. His hypothesis was that, by selecting a behavioral trait, he could also influence the phenotype of subsequent generations, making them more domestic in appearance. Over the next 40 years, he succeeded in producing foxes with traits that were never directly selected for, including piebald coats floppy ears, upturned tails, shortened snouts, and shifts in developmental timing. In the 1980s, a researcher used a set of behavioral, cognitive, and visible phenotypic markers, such as coat colour, to produce domesticated fallow deer within a few generations. Similar results for tameness and fear have been found for mink and Japanese quail. In addition to demonstrating that domestic phenotypic traits could arise through selection for a behavioral trait, and domestic behavioral traits could arise through the selection for a phenotypic trait, these experiments provided a mechanism to explain how the animal domestication process could have begun without deliberate human forethought and action.
The genetic difference between domestic and wild populations can be framed within two considerations. The first distinguishes between domestication traits that are presumed to have been essential at the early stages of domestication, and improvement traits that have appeared since the split between wild and domestic populations. Domestication traits are generally fixed within all domesticates and were selected during the initial episode of domestication, whereas improvement traits are present only in a proportion of domesticates, though they may be fixed in individual breeds or regional populations. A second issue is whether traits associated with the domestication syndrome resulted from a relaxation of selection as animals exited the wild environment or from positive selection resulting from intentional and unintentional human preference. Some recent genomic studies on the genetic basis of traits associated with the domestication syndrome have shed light on both of these issues. A study published in 2016 suggested that there have been negative genetic consequences of the domestication process as well, that enrichment of disease-related gene variants accompanied positively selected traits.
In 2010, a study identified 51 regions of the dog genome that were associated with phenotypic variation among breeds in 57 traits studied, which included body, cranial, dental, and long bone shape and size. There were 3 quantitative trait loci that explained most of the phenotypic variation. Indicators of recent selection were shown by many of the 51 genomic regions that were associated with traits that define a breed, which include body size, coat characteristics, and ear floppiness. Geneticists have identified more than 300 genetic loci and 150 genes associated with coat color variability. Knowing the mutations associated with different colors has allowed the correlation between the timing of the appearance of variable coat colors in horses with the timing of their domestication. Other studies have shown how human-induced selection is responsible for the allelic variation in pigs. Together, these insights suggest that, although natural selection has kept variation to a minimum before domestication, humans have actively selected for novel coat colors as soon as they appeared in managed populations.
In 2015, a study looked at over 100 pig genome sequences to ascertain their process of domestication. A model that fitted the data included admixture with a now extinct ghost population of wild pigs during the Pleistocene. The study also found that despite back-crossing with wild pigs, the genomes of domestic pigs have strong signatures of selection at genetic loci that affect behavior and morphology. The study concluded that human selection for domestic traits likely counteracted the homogenizing effect of gene flow from wild boars and created domestication islands in the genome. The same process may also apply to other domesticated animals.
In 2014, a whole genome study of the DNA differences between wolves and dogs found that dogs did not show a reduced fear response but did show greater synaptic plasticity. Synaptic plasticity is widely believed to be the cellular correlate of learning and memory, and this change may have altered the learning and memory abilities of dogs in comparison to wolves.
Unlike other domestic species which were primarily selected for production-related traits, dogs were initially selected for their behaviors. In 2016, a study found that there were only 11 fixed genes that showed variation between wolves and dogs. These gene variations were unlikely to have been the result of natural evolution, and indicate selection on both morphology and behavior during dog domestication. There was evidence of selection during dog domestication of genes that affect the adrenaline and noradrenaline biosynthesis pathway. These genes are involved in the synthesis, transport and degradation of a variety of neurotransmitters, particularly the catecholamines, which include dopamine and noradrenaline. Recurrent selection on this pathway and its role in emotional processing and the fight-or-flight response suggests that the behavioral changes we see in dogs compared to wolves may be due to changes in this pathway, leading to tameness and an emotional processing ability. Dogs generally show reduced fear and aggression compared to wolves. Some of these genes have been associated with aggression in some dog breeds, indicating their importance in both the initial domestication and then later in breed formation.
In 2018, a study identified 429 genes that differed between modern dogs and modern wolves. As the differences in these genes could also be found in ancient dog fossils, these were regarded as being the result of the initial domestication and not from recent breed formation. These genes are linked to neural crest and central nervous system development. These genes affect embryogenesis and can confer tameness, smaller jaws, floppy ears, and diminished craniofacial development, which distinguish domesticated dogs from wolves and are considered to reflect domestication syndrome. The study proposes that domestication syndrome is caused by alterations in the migration or activity of neural crest cells during their development. The study concluded that during early dog domestication, the initial selection was for behavior. This trait is influenced by those genes which act in the neural crest, which led to the phenotypes observed in modern dogs.
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. In a study in 2014, the data indicated that the wolves and dingo had just two copies of the gene and the Siberian Husky that is associated with hunter-gatherers had just three or four 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. A later study indicated that because most dogs had a high copy number of the AMY2B gene but the arctic breeds and the dingo did not, that the dog's dietary change may not have been caused by initial domestication but by the subsequent spread of agriculture to most - but not all - regions of the planet.
In 2016, a study of the dog genome compared to the wolf genome looked for genes that showed signs of having undergone positive selection. The study identified genes relating to brain function and behavior, and to lipid metabolism. This ability to process lipids indicates a dietary target of selection that was important when proto-dogs hunted and fed alongside hunter-gatherers. The evolution of the dietary metabolism genes may have helped process the increased lipid content of early dog diets as they scavenged on the remains of carcasses left by hunter-gatherers. Prey capture rates may have increased in comparison to wolves and with it the amount of lipid consumed by the assisting proto-dogs. A unique dietary selection pressure may have evolved both from the amount consumed, and the shifting composition of, tissues that were available to proto-dogs once humans had removed the most desirable parts of the carcass for themselves. A study of the mammal biomass during modern human expansion into the northern Mammoth steppe found that it had occurred under conditions of unlimited resources, and that many of the animals were killed with only a small part consumed or left unused.
- See further: Phenotypic plasticity
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.":92
- See further: Dog learning by inference
A dog's cranium is 15% smaller than an equally heavy wolf's, and the dog is less aggressive and more playful. Other species pairs show similar differences. Bonobos, like chimpanzees, are a close genetic cousin to humans, but unlike the chimpanzees, bonobos are not aggressive and do not participate in lethal inter-group 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. In other examples, the guinea pig's cranium is 13% smaller than its wild cousin the cavy, 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.:104 In the farm fox experiment, humans selectively bred foxes against aggression, causing 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 favors 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.:114 The more docile animals have been found to have less testosterone than their more aggressive counterparts, and testosterone controls aggression and brain size. One researcher has argued that in becoming more social, we humans have developed a smaller brain than those of humans 20,000 years ago.
Dog and human convergent evolution
As a result of the domestication process there is also evidence of convergent evolution having occurred between dogs and humans. The history of the two is forever intertwined. Dogs suffer from the same diseases as humans, which include cancer, diabetes, heart disease, and neurological disorders. The underlying disease pathology is similar to humans, as is their responses and outcomes to treatment.
There are patterns of genes which are related by their function and these patterns can be found in both dogs and humans. This fact can be used to study the coevolution of gene function. Dogs accompanied humans when they first migrated into new environments. Both dogs and humans have adapted to different environmental conditions, with their genomes showing parallel evolution. These include adaptation to high altitude, low oxygen hypoxia conditions, and genes that play a role in digestion, metabolism, neurological processes, and some related to cancer. It can be inferred from those genes which act on the serotonin system in the brain that these have given rise to less aggressive behavior when living in a crowded environment.
In 2007, a study found that dog domestication was accompanied by selection at three genes with key roles in starch digestion: AMY2B, MGAMand SGLT1, and was a striking case of parallel evolution when coping with an increasingly starch-rich diet caused similar adaptive responses in dogs and humans.
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 a solution to the problem of moving through the water. What has been found between dogs and humans 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.: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 phylogenetically, 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.
The pointing gesture is a human-specific signal, is referential in its nature, and is a foundation building-block of human communication. Human infants acquire it weeks before the first spoken word. 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 demonstrating a similar level of performance as 2-year-old children can be explained as a joint outcome of their evolutionary history as well as their socialization in a human environment.
Later studies support coevolution in that dogs can discriminate the emotional expressions of human faces, and that most people can tell from a bark whether a dog is alone, being approached by a stranger, playing, or being aggressive, and can tell from a growl how big the dog is.
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. One observer has stated, "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....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 they 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 in ways, we must assume, they could never have imagined. Powerful emotions were in play that many observers today refer to as love – boundless, unquestioning love."
Human adoption of some wolf behaviors
|“||... 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? ...||”|
|— Wolfgang Schleidt|
In 2002, a study proposed 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. 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 led 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. One of the keys to recent human survival has been the forming of partnerships. Strong bonds exist between same-sex wolves, dogs and humans and these bonds are stronger than exist 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.
In 2003, a study compared the behavior and ethics of chimpanzees, wolves and humans. Cooperation among humans' closest genetic relative is limited to occasional hunting episodes or the persecution of a competitor for personal advantage, which had to be tempered if humans were to become domesticated. The closest approximation to human morality that can be found in nature is that of the gray wolf, Canis lupus. Wolves are among the most gregarious and cooperative of animals on the planet, and their 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 rivaled only by that of human societies. Similar forms of cooperation are observed in two closely related canids, the African wild 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. Social sharing within families may be a trait that early humans learned from wolves, and with wolves digging dens long before humans constructed huts it is not clear who domesticated whom.
On the mammoth steppe the wolf's ability to hunt in packs, to share risk fairly among pack members, and to cooperate moved them to the top of the food chain above lions, hyenas and bears. Some 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 study proposed that during the Last Glacial Maximum, some of our ancestors teamed up with those pastoralist wolves and learned their techniques. Many of our ancestors remained gatherers and scavengers, or specialized as fish-hunters, hunter-gatherers, and 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, and this group of humans was to become the first herders and this group of wolves was to become the first dogs.
The dog was the first species and the only large carnivore to have been domesticated. Over the past 200 years, dogs have undergone rapid phenotypic change and were formed into today's modern dog breeds due to artificial selection imposed by humans. These breeds can vary in size and weight from a 0.46 kg (1.0 lb) teacup poodle to a 90 kg (200 lb) giant mastiff. The skull, body, and limb proportions vary significantly between breeds, with dogs displaying more phenotypic diversity than can be found within the entire order of carnivores. Some breeds demonstrate outstanding skills in herding, retrieving, scent detection, and guarding, which demonstrates the functional and behavioral diversity of dogs. There have been major advances in understanding the genes that gave rise to the phenotypic traits of dogs. The first dogs were certainly wolflike, however the phenotypic changes that coincided with the dog–wolf genetic divergence are not known.
In 1914, on the eve of the First World War, two human skeletons were discovered during basalt quarrying at Oberkassel, Bonn in Germany. With them were found a right mandible of a "wolf" and other animal bones. After the end of the First World War, in 1919 a full study was made of these remains. The mandible was recorded as "Canis lupus, the wolf" and some of the other animal bones were assigned to it. The remains were then stored and forgotten for fifty years. In the late 1970s there was renewed interest in the Oberkassel remains and the mandible was re-examined and reclassified as belonging to a domesticated dog. The mitochondrial DNA sequence of the mandible was matched to Canis lupus familiaris - dog, and confirms that the Oberkassel dog is a direct ancestor of today's dogs. The bodies were dated to 14,223 YBP. This implies that in Western Europe there were morphologically and genetically "modern" dogs in existence around 14,500 years ago.
Later studies assigned more of the other animal bones to the dog until most of a skeleton could be assembled. The humans were a man aged 40 years and a woman aged 25 years. All three skeletal remains were found covered with large 20 cm thick basalt blocks and were sprayed with red hematite powder. The consensus is that a dog was buried along with two humans. A tooth belonging to a smaller and older dog was also identified but it had not been sprayed with red powder. The cause of the death of the two humans is not known. A pathology study of the dog remains suggests that it had died young after suffering from canine distemper between ages 19 and 23 weeks. The dog could not have survived during this period without intensive human care. During this period the dog was of no utilitarian use to humans, and suggests the existence of emotional or symbolic ties between these humans and this dog. In conclusion, near the end of the Late Pleistocene at least some humans regarded dogs not just materialistically, but had developed emotional and caring bonds for their dogs.
First dogs as a hunting technology
During the Upper Paleolithic (50,000-10,000 YBP), the increase in human population density, advances in blade and hunting technology, and climate change may have altered prey densities and made scavenging crucial to the survival of some wolf populations. Adaptations to scavenging such as tameness, small body size, and a decreased age of reproduction would reduce their hunting efficiency further, eventually leading to obligated scavenging. Whether these earliest dogs were simply human-commensal scavengers or they played some role as companions or hunters that hastened their spread is unknown.
Researchers have proposed that in the past a hunting partnership existed between humans and dogs that was the basis for dog domestication. Petroglyph rock art dating to 8,000 YBP at the sites of Shuwaymis and Jubbah, in northwestern Saudi Arabia, depict large numbers of dogs participating in hunting scenes with some being controlled on leashes. The transition from the Late Pleistocene into the early Holocene was marked by climatic change from cold and dry to warmer, wetter conditions and rapid shifts in flora and fauna, with much of the open habitat of large herbivores being replaced by forests. In the early Holocene, it is proposed that along with changes in arrow-head technology that hunting dogs were used by hunters to track and retrieve wounded game in thick forests. The dog's ability to chase, track, sniff out and hold prey can significantly increase the success of hunters in forests, where human senses and location skills are not as sharp as in more open habitats. Dogs are still used for hunting in forests today.
Dogs enter North America from Siberia
In North America, the earliest dog remains were found in Illinois and radiocarbon dating indicates 9,900 YBP. These include three isolated burials at the Koster Site near the lower Illinois River in Greene County, and one burial 35 km away at the Stilwell II site in Pike County. These dogs were medium-sized adults around 50 cm (20 in) in height and around 17 kilograms (37 lb) in weight, with very active lifestyles and varied morphologies. Isotope analysis can be used to identify some chemical elements, allowing researchers to make inferences about the diet of a species. An isotope analysis of bone collagen indicates a diet consisting largely of freshwater fish. Similar dog burials across Eurasia are thought to be due to the dog’s importance in hunting to people who were trying to adapt to the changing environments and prey species during the Pleistocene-Holocene transition. In these places, the dog had gained an elevated social status. 
In 2018, a study compared sequences of North American dog fossils with Siberian dog fossils and modern dogs. The nearest relative to the North American fossils was a 9,000 YBP fossil discovered on Zhokhov Island, arctic north-eastern Siberia, which was connected to the mainland at that time. The study inferred from mDNA that all of the North American dogs shared a common ancestor dated 14,600 YBP, and this ancestor had diverged along with the ancestor of the Zhokhov dog from their common ancestor 15,600 YBP. The timing of the Koster dogs shows that dogs entered North America from Siberia 4,500 years after humans did, were isolated for the next 9,000 years, and after contact with Europeans these no longer exist because they were replaced by Eurasian dogs. The pre-contact dogs exhibit a unique genetic signature that is now gone, with nDNA indicating that their nearest genetic relatives today are the arctic breed dogs - Alaskan malamutes, Greenland dogs, and Alaskan huskies and Siberian huskies.
First dog breeds developed in Siberia
In 2017, a study showed that 9,000 YBP the domestic dog was present at what is now Zhokhov Island. The dogs were selectively bred as either sled dogs or as hunting dogs, which implies that a sled dog standard and a hunting dog standard existed at that time. The optimal maximum size for a sled dog is 20–25 kg based on thermo-regulation, and the ancient sled dogs were between 16–25 kg. The same standard has been found in the remains of sled dogs from this region 2,000 YBP and in the modern Siberian husky breed standard. Other dogs were more massive at 30 kg and appear to be dogs that had been crossed with wolves and used for polar bear hunting. At death, the heads of the dogs had been carefully separated from their bodies by humans, probably for ceremonial reasons.
The study proposes that after having diverged from the common ancestor along with the grey wolf, the evolution of Canis familiaris proceeded in three stages. The first was natural selection based on feeding behavior within the ecological niche that had been formed through human activity. The second was artificial selection based on tamability. The third was directed selection based on forming breeds that possessed qualities to help with specific tasks within the human economy. The process commenced 40,000-30,000 YBP with its speed increasing with each stage until domestication became complete.
Dogs enter Japan
In Japan, temperate deciduous forests rapidly spread onto the main island of Honshu and caused an adaption away from hunting megafauna (Naumann's elephant and Yabe's giant deer) to hunting the quicker sika deer and wild boar in dense forest. With this came a change in hunting technology, including a shift to smaller, triangular points for arrows. A study of the Jōmon people that lived on the Pacific coast of Honshu during the early Holocene shows that they were conducting individual dog burials and were probably using dogs as tools for hunting sika deer and wild boar, as hunters in Japan still do today.
Hunting dogs make major contributions to forager societies and the ethnographic record shows them being given proper names, treated as family members, and considered separate to other types of dogs. This special treatment includes separate burials with markers and grave-goods, with those that were exceptional hunters or that were killed on the hunt often venerated. A dog's value as a hunting partner gives them status as a living weapon and the most skilled elevated to taking on a "personhood", with their social position in life and in death similar to that of the skilled hunters.
Intentional dog burials together with ungulate hunting is also found in other early Holocene deciduous forest forager societies in Europe and North America, indicating that across the Holarctic temperate zone hunting dogs were a widespread adaptation to forest ungulate hunting.
- Larson, G.; Bradley, D.G. (2014). "How Much Is That in Dog Years? The Advent of Canine Population Genomics". PLOS Genetics. 10 (1): e1004093. doi:10.1371/journal.pgen.1004093. PMC 3894154. PMID 24453989.
- Freedman, Adam H; Wayne, Robert K (2017). "Deciphering the Origin of Dogs: From Fossils to Genomes". Annual Review of Animal Biosciences. 5: 281–307. doi:10.1146/annurev-animal-022114-110937. PMID 27912242.
- Irving-Pease, Evan K.; Ryan, Hannah; Jamieson, Alexandra; Dimopoulos, Evangelos A.; Larson, Greger; Frantz, Laurent A. F. (2018). "Paleogenomics of Animal Domestication". In Lindqvist, C.; Rajora, O. (eds.). Paleogenomics. Population Genomics. Springer, Cham. pp. 225–272. doi:10.1007/13836_2018_55. ISBN 978-3-030-04752-8.
- Thalmann, Olaf; Perri, Angela R. (2018). "Paleogenomic Inferences of Dog Domestication". In Lindqvist, C.; Rajora, O. (eds.). Paleogenomics. Population Genomics. Springer, Cham. pp. 273–306. doi:10.1007/13836_2018_27. ISBN 978-3-030-04752-8.
- Machugh, David E.; Larson, Greger; Orlando, Ludovic (2016). "Taming the Past: Ancient DNA and the Study of Animal Domestication". Annual Review of Animal Biosciences. 5: 329–351. doi:10.1146/annurev-animal-022516-022747. PMID 27813680.
- Thalmann, O; Shapiro, B; Cui, P; Schuenemann, V. J; Sawyer, S. K; Greenfield, D. L; Germonpre, M. B; Sablin, M. V; Lopez-Giraldez, 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; Paabo, S; et al. (2013). "Complete Mitochondrial Genomes of Ancient Canids Suggest a European Origin of Domestic Dogs". Science. 342 (6160): 871–4. Bibcode:2013Sci...342..871T. doi:10.1126/science.1243650. PMID 24233726.
- Shannon, L (2015). "Genetic structure in village dogs reveals a Central Asian domestication origin". Proceedings of the National Academy of Sciences. 112 (44): 13639–44. Bibcode:2015PNAS..11213639S. doi:10.1073/pnas.1516215112. PMC 4640804. PMID 26483491.
- Wang, G (2015). "Out of southern East Asia: the natural history of domestic dogs across the world". Cell Research. 26 (1): 21–33. doi:10.1038/cr.2015.147. PMC 4816135. PMID 26667385.
- R.M. Nowak (2003). "Chapter 9 - Wolf evolution and taxonomy". In Mech, L. David; Boitani, Luigi (eds.). Wolves: Behaviour, Ecology and Conservation. University of Chicago Press. pp. 239–258. ISBN 978-0-226-51696-7. page 239
- Pierotti & Fogg 2017, pp. 5–6
- Derr 2011, pp. 40
- Cooper, A. (2015). "Abrupt warming events drove Late Pleistocene Holarctic megafaunal turnover". Science. 349 (6248): 602–6. Bibcode:2015Sci...349..602C. doi:10.1126/science.aac4315. PMID 26250679.
- Fan, Zhenxin; Silva, Pedro; Gronau, Ilan; Wang, Shuoguo; Armero, Aitor Serres; Schweizer, Rena M.; Ramirez, Oscar; Pollinger, John; Galaverni, Marco; Ortega Del-Vecchyo, Diego; Du, Lianming; Zhang, Wenping; Zhang, Zhihe; Xing, Jinchuan; Vilà, Carles; Marques-Bonet, Tomas; Godinho, Raquel; Yue, Bisong; Wayne, Robert K. (2016). "Worldwide patterns of genomic variation and admixture in gray wolves". Genome Research. 26 (2): 163–73. doi:10.1101/gr.197517.115. PMC 4728369. PMID 26680994.
- Skoglund, Pontus; Ersmark, Erik; Palkopoulou, Eleftheria; Dalén, Love (2015). "Ancient Wolf Genome Reveals an Early Divergence of Domestic Dog Ancestors and Admixture into High-Latitude Breeds". Current Biology. 25 (11): 1515–1519. doi:10.1016/j.cub.2015.04.019. PMID 26004765.
- Lindblad-Toh, K.; Wade, C. M.; Mikkelsen, T. S.; Karlsson, E. K.; Jaffe, D. B.; Kamal, M.; Clamp, M.; Chang, J. L.; Kulbokas, E. J.; Zody, M. C.; Mauceli, E.; Xie, X.; Breen, M.; Wayne, R. K.; Ostrander, E. A.; Ponting, C. P.; Galibert, F.; Smith, D. R.; Dejong, P. J.; Kirkness, E.; Alvarez, P.; Biagi, T.; Brockman, W.; Butler, J.; Chin, C. W.; Cook, A.; Cuff, J.; Daly, M. J.; Decaprio, D.; et al. (2005). "Genome sequence, comparative analysis and haplotype structure of the domestic dog". Nature. 438 (7069): 803–819. Bibcode:2005Natur.438..803L. doi:10.1038/nature04338. PMID 16341006.
- Lee, E. (2015). "Ancient DNA analysis of the oldest canid species from the Siberian Arctic and genetic contribution to the domestic dog". PLoS ONE. 10 (5): e0125759. Bibcode:2015PLoSO..1025759L. doi:10.1371/journal.pone.0125759. PMC 4446326. PMID 26018528.
- Irizarry, Kristopher J. L.; Vasconcelos, Elton J. R. (2018). "Population Genomics of Domestication and Breed Development in Canines in the Context of Cognitive, Social, Behavioral, and Disease Traits". In Rajora, O. (ed.). Population Genomics. pp. 755–806. doi:10.1007/13836_2018_43. ISBN 978-3-030-04587-6.
- vonHoldt, B. (2010). "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.
- Frantz, L. A. F.; Mullin, V. E.; Pionnier-Capitan, M.; Lebrasseur, O.; Ollivier, M.; Perri, A.; Linderholm, A.; Mattiangeli, V.; Teasdale, M. D.; Dimopoulos, E. A.; Tresset, A.; Duffraisse, M.; McCormick, F.; Bartosiewicz, L.; Gal, E.; Nyerges, E. A.; Sablin, M. V.; Brehard, S.; Mashkour, M.; b l Escu, A.; Gillet, B.; Hughes, S.; Chassaing, O.; Hitte, C.; Vigne, J.-D.; Dobney, K.; Hanni, C.; Bradley, D. G.; Larson, G. (2016). "Genomic and archaeological evidence suggest a dual origin of domestic dogs". Science. 352 (6290): 1228–31. Bibcode:2016Sci...352.1228F. doi:10.1126/science.aaf3161. PMID 27257259.
- Leonard, J. A.; Wayne, R. K.; Wheeler, J; Valadez, R; Guillén, S; Vilà, C (2002). "Ancient DNA Evidence for Old World Origin of New World Dogs". Science. 298 (5598): 1613–6. Bibcode:2002Sci...298.1613L. doi:10.1126/science.1076980. PMID 12446908.
- Freedman, Adam H.; Lohmueller, Kirk E.; Wayne, Robert K. (2016). "Evolutionary History, Selective Sweeps, and Deleterious Variation in the Dog". Annual Review of Ecology, Evolution, and Systematics. 47: 73–96. doi:10.1146/annurev-ecolsys-121415-032155.
- Malmström, Helena; Vilà, Carles; Gilbert, M; Storå, Jan; Willerslev, Eske; Holmlund, Gunilla; Götherström, Anders (2008). "Barking up the wrong tree: Modern northern European dogs fail to explain their origin". BMC Evolutionary Biology. 8: 71. doi:10.1186/1471-2148-8-71. PMC 2288593. PMID 18307773.
- Larson G (2012). "Rethinking dog domestication by integrating genetics, archeology, and biogeography". PNAS. 109 (23): 8878–8883. Bibcode:2012PNAS..109.8878L. doi:10.1073/pnas.1203005109. PMC 3384140. PMID 22615366.
- Frantz, L. (2015). "Evidence of long-term gene flow and selection during domestication from analyses of Eurasian wild and domestic pig genomes". Nature Genetics. 47 (10): 1141–1148. doi:10.1038/ng.3394. PMID 26323058.
- vonHoldt, Bridgett M.; Driscoll, Carlos A. (2016). "3-Origins of the dog:Genetic insights into dog domestication". In James Serpell (ed.). The Domestic Dog: Its Evolution, Behavior and Interactions with People (2 ed.). Cambridge University Press. pp. 22–41. ISBN 978-1-107-02414-4.
- Miklosi, Adam (2018). "1-Evolution & Ecology". The Dog: A Natural History. Princeton University Press. pp. 13–39. ISBN 978-0-691-17693-2.
- Melissa Chan (2016). The Mysterious History Behind Humanity's Love of Dogs (Interview with Greger Larson). Time.
- Schleidt, Wolfgang M.; Shalter, Michael D. (2018). "Dogs and Mankind: Coevolution on the Move – an Update". Human Ethology Bulletin. 33: 15–38. doi:10.22330/heb/331/015-038.
- Botigué, Laura R.; Song, Shiya; Scheu, Amelie; Gopalan, Shyamalika; Pendleton, Amanda L.; Oetjens, Matthew; Taravella, Angela M.; Seregély, Timo; Zeeb-Lanz, Andrea; Arbogast, Rose-Marie; Bobo, Dean; Daly, Kevin; Unterländer, Martina; Burger, Joachim; Kidd, Jeffrey M.; Veeramah, Krishna R. (2017). "Ancient European dog genomes reveal continuity since the Early Neolithic". Nature Communications. 8: 16082. Bibcode:2017NatCo...816082B. doi:10.1038/ncomms16082. PMC 5520058. PMID 28719574.
- Ollivier, Morgane; Tresset, Anne; Frantz, Laurent A. F.; Bréhard, Stéphanie; Bălăşescu, Adrian; Mashkour, Marjan; Boroneanţ, Adina; Pionnier-Capitan, Maud; Lebrasseur, Ophélie; Arbogast, Rose-Marie; Bartosiewicz, László; Debue, Karyne; Rabinovich, Rivka; Sablin, Mikhail V.; Larson, Greger; Hänni, Catherine; Hitte, Christophe; Vigne, Jean-Denis (2018). "Dogs accompanied humans during the Neolithic expansion into Europe". Biology Letters. 14 (10): 20180286. doi:10.1098/rsbl.2018.0286. PMC 6227856. PMID 30333260.
- Oetjens, Matthew T.; Martin, Axel; Veeramah, Krishna R.; Kidd, Jeffrey M. (2018). "Analysis of the canid Y-chromosome phylogeny using short-read sequencing data reveals the presence of distinct haplogroups among Neolithic European dogs". BMC Genomics. 19 (1): 350. doi:10.1186/s12864-018-4749-z. PMC 5946424. PMID 29747566.
- Irving-Pease, Evan K; Frantz, Laurent A.F; Sykes, Naomi; Callou, Cécile; Larson, Greger (2018). "Rabbits and the Specious Origins of Domestication". Trends in Ecology & Evolution. 33 (3): 149–152. doi:10.1016/j.tree.2017.12.009. PMID 29454669.
- Ed Yong (2016). "A New Origin Story for Dogs - Interview with Greger Larson". The Atlantic Monthly Group.
- Grimm, David (2015). "Feature: Solving the mystery of dog domestication". Science. doi:10.1126/science.aab2477. quoting Greger Larson
- Ostrander, Elaine A.; Wang, Guo-Dong; Larson, Greger; Vonholdt, Bridgett M.; Davis, Brian W.; Jagannathan, Vidyha; Hitte, Christophe; Wayne, Robert K.; Zhang, Ya-Ping (2019). "Dog10K: An international sequencing effort to advance studies of canine domestication, phenotypes, and health". National Science Review. doi:10.1093/nsr/nwz049.
- Morey, Darcy F.; Jeger, Rujana (2016). "From wolf to dog: Late Pleistocene ecological dynamics, altered trophic strategies, and shifting human perceptions". Historical Biology. 29 (7): 895–903. doi:10.1080/08912963.2016.1262854.
- Duleba, Anna; Skonieczna, Katarzyna; Bogdanowicz, Wiesław; Malyarchuk, Boris; Grzybowski, Tomasz (2015). "Complete mitochondrial genome database and standardized classification system for Canis lupus familiaris". Forensic Science International: Genetics. 19: 123–129. doi:10.1016/j.fsigen.2015.06.014. PMID 26218982.
- Pierotti & Fogg 2017, pp. 192–193
- Jans, N. (2014). A wolf called Romeo. Houghton Mifflin Harcourt. ISBN 978-0547858197.
- Derr 2011, pp. 85–98
- Wang, Xiaoming; Tedford, Richard H.; Dogs: Their Fossil Relatives and Evolutionary History. New York: Columbia University Press, 2008. pp. 166
- Schleidt, W. (2003). "Co-evolution of humans and canids: An alternative view of dog domestication: Homo homini lupus?" (PDF). Evolution and Cognition. 9 (1): 57–72.
- Frans de Waal (2006). Primates and Philosophers: How Morality Evolved. Princeton University Press. p. 3.
- Vila, C. (1997). "Multiple and ancient origins of the domestic dog". Science. 276 (5319): 1687–9. doi:10.1126/science.276.5319.1687. PMID 9180076.
- Olsen, S. J. (1985). Origins of the domestic dog: the fossil record. Univ. of Arizona Press, Tucson, US. pp. 88–89.
- Zeder MA (2012). "The domestication of animals". Journal of Anthropological Research. 68 (2): 161–190. doi:10.3998/jar.0521004.0068.201.
- Lyudmila N. Trut (1999). "Early Canid Domestication: The Farm-Fox Experiment" (PDF). American Scientist. 87 (March–April): 160–169. doi:10.1511/1999.2.160. Archived from the original (PDF) on February 15, 2010. Retrieved January 12, 2016.
- Morey Darcy F (1992). "Size, shape, and development in the evolution of the domestic dog". Journal of Archaeological Science. 19 (2): 181–204. doi:10.1016/0305-4403(92)90049-9.
- Turnbull Priscilla F.; Reed Charles A. (1974). "The fauna from the terminal Pleistocene of Palegawra Cave". Fieldiana: Anthropology. 63: 81–146.
- Musiani M, Leonard JA, Cluff H, Gates CC, Mariani S, et al. (2007). "Differentiation of tundra/taiga and boreal coniferous forest wolves: genetics, coat colour and association with migratory caribou". Mol. Ecol. 16 (19): 4149–70. doi:10.1111/j.1365-294x.2007.03458.x. PMID 17725575.
- Leonard, J. (2007). "Megafaunal extinctions and the disappearance of a specialized wolf ecomorph". Current Biology. 17 (13): 1146–50. doi:10.1016/j.cub.2007.05.072. PMID 17583509.
- Wolpert, S. (2013), "Dogs likely originated in Europe more than 18,000 years ago, UCLA biologists report", UCLA News Room, retrieved December 10, 2014
- Arctic Wolf: The High Arctic by Laura DeLallo. Bearport Publishing, New York 2011
- Arctic wildlife in a warming world by Michael Becker. BBC Two, 2014.
- Ellesmere Island Journal & Field Notes by Henry Beston 2006. International Wolf Centre.
- Arctic Wolves and Their Prey by L. David Mech. National Ocean and Atmospheric Administration, Pacific Marine Environment Laboratory, Actic Zone. 2004
- Pang, J. (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. 26 (12): 2849–64. doi:10.1093/molbev/msp195. PMC 2775109. PMID 19723671.
- Freedman, A. (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.
- Ardalan, A (2011). "Comprehensive study of mtDNA among Southwest Asian dogs contradicts independent domestication of wolf, but implies dog–wolf hybridization". Ecology and Evolution. 1 (3): 373–385. doi:10.1002/ece3.35. PMC 3287314. PMID 22393507.
- Klutsch, C.F. (2010). "Regional occurrence, high frequency but low diversity of mitochondrial DNA haplogroup d1 suggests a recent dog-wolf hybridization in Scandinavia". Journal of Veterinary Behavior: Clinical Applications and Research. 6 (1): 100–103. doi:10.1016/j.jveb.2010.08.035. PMC 3040290. PMID 20497152.
- Ishiguro, N (2009). "Mitochondrial DNA Analysis of the Japanese Wolf (Canis Lupus Hodophilax Temminck, 1839) and Comparison with Representative Wolf and Domestic Dog Haplotypes". Zoological Science. 26 (11): 765–770. doi:10.2108/zsj.26.765. PMID 19877836.
- Bjornerfeldt, S (2006). "Relaxation of selective constraint on dog mitochondrial DNA followed domestication". Genome Research. 16 (8): 990–994. doi:10.1101/gr.5117706. PMC 1524871. PMID 16809672.
- Savolainen, P. (2002). "Genetic evidence for an East Asian origin of domestic dogs". Science. 298 (5598): 1610–3. Bibcode:2002Sci...298.1610S. doi:10.1126/science.1073906. PMID 12446907.
- Pilot, Małgorzata; Greco, Claudia; Vonholdt, Bridgett M; Randi, Ettore; Jędrzejewski, Włodzimierz; Sidorovich, Vadim E; Konopiński, Maciej K; Ostrander, Elaine A; Wayne, Robert K (2018). "Widespread, long-term admixture between grey wolves and domestic dogs across Eurasia and its implications for the conservation status of hybrids". Evolutionary Applications. 11 (5): 662–680. doi:10.1111/eva.12595. PMC 5978975. PMID 29875809.
- Schweizer, Rena M; Durvasula, Arun; Smith, Joel; Vohr, Samuel H; Stahler, Daniel R; Galaverni, Marco; Thalmann, Olaf; Smith, Douglas W; Randi, Ettore; Ostrander, Elaine A; Green, Richard E; Lohmueller, Kirk E; Novembre, John; Wayne, Robert K (2018). "Natural Selection and Origin of a Melanistic Allele in North American Gray Wolves". Molecular Biology and Evolution. 35 (5): 1190–1209. doi:10.1093/molbev/msy031. PMID 29688543.
- Darwin, Charles (1868). "Chapter 1: Domestic Dogs and Cats". The Variation of Animals and Plants under Domestication. Vol. 1. John Murray, London.
- Larson, G.; Piperno, D. R.; Allaby, R. G.; Purugganan, M. D.; Andersson, L.; Arroyo-Kalin, M.; Barton, L.; Climer Vigueira, C.; Denham, T.; Dobney, K.; Doust, A. N.; Gepts, P.; Gilbert, M. T. P.; Gremillion, K. J.; Lucas, L.; Lukens, L.; Marshall, F. B.; Olsen, K. M.; Pires, J. C.; Richerson, P. J.; Rubio De Casas, R.; Sanjur, O. I.; Thomas, M. G.; Fuller, D. Q. (2014). "Current perspectives and the future of domestication studies". Proceedings of the National Academy of Sciences. 111 (17): 6139–6146. Bibcode:2014PNAS..111.6139L. doi:10.1073/pnas.1323964111. PMC 4035915. PMID 24757054.
- Larson, G (2013). "A population genetics view of animal domestication" (PDF).
- Trut, L.; et al. (2009). "Animal evolution during domestication: the domesticated fox as a model". BioEssays. 31 (3): 349–360. doi:10.1002/bies.200800070. PMC 2763232. PMID 19260016.
- Hemmer H (2005). "Neumuhle-Riswicker Hirsche: Erste planma¨ßige Zucht einer neuen Nutztierform". Naturwissenschaftliche Rundschau. 58: 255–261.
- Malmkvist, Jen s; Hansen, Steffen W. (2002). "Generalization of fear in farm mink, Mustela vison, genetically selected for behaviour towards humans" (PDF). Animal Behaviour. 64 (3): 487–501. doi:10.1006/anbe.2002.3058.
- Jones, R.Bryan; Satterlee, Daniel G.; Marks, Henry L. (1997). "Fear-related behaviour in Japanese quail divergently selected for body weight". Applied Animal Behaviour Science. 52 (1–2): 87–98. doi:10.1016/S0168-1591(96)01146-X.
- Olsen KM, Wendel JF (2013). "A bountiful harvest: genomic insights into crop domestication phenotypes". Annu. Rev. Plant Biol. 64: 47–70. doi:10.1146/annurev-arplant-050312-120048. PMID 23451788.
- Doust, A. N.; Lukens, L.; Olsen, K. M.; Mauro-Herrera, M.; Meyer, A.; Rogers, K. (2014). "Beyond the single gene: How epistasis and gene-by-environment effects influence crop domestication". Proceedings of the National Academy of Sciences. 111 (17): 6178–6183. Bibcode:2014PNAS..111.6178D. doi:10.1073/pnas.1308940110. PMC 4035984. PMID 24753598.
- Meyer, Rachel S.; Purugganan, Michael D. (2013). "Evolution of crop species: Genetics of domestication and diversification". Nature Reviews Genetics. 14 (12): 840–52. doi:10.1038/nrg3605. PMID 24240513.
- Marsden, Clare D.; Vecchyo, Diego Ortega-Del; O'Brien, Dennis P.; et al. (5 January 2016). "Bottlenecks and selective sweeps during domestication have increased deleterious genetic variation in dogs". Proceedings of the National Academy of Sciences of the United States of America. 113 (1): 152–7. Bibcode:2016PNAS..113..152M. doi:10.1073/pnas.1512501113. PMC 4711842. PMID 26699508.
- Boyko, Adam R.; Quignon, Pascale; Li, Lin; Schoenebeck, Jeffrey J.; Degenhardt, Jeremiah D.; Lohmueller, Kirk E.; Zhao, Keyan; Brisbin, Abra; Parker, Heidi G.; Vonholdt, Bridgett M.; Cargill, Michele; Auton, Adam; Reynolds, Andy; Elkahloun, Abdel G.; Castelhano, Marta; Mosher, Dana S.; Sutter, Nathan B.; Johnson, Gary S.; Novembre, John; Hubisz, Melissa J.; Siepel, Adam; Wayne, Robert K.; Bustamante, Carlos D.; Ostrander, Elaine A. (2010). "A Simple Genetic Architecture Underlies Morphological Variation in Dogs". PLoS Biology. 8 (8): e1000451. doi:10.1371/journal.pbio.1000451. PMC 2919785. PMID 20711490. This article incorporates text from this source, which is in the public domain.
- Cieslak, M. et al. (2011) Colours of domestication. Biol. Rev. 86, 885–899
- Ludwig A.; et al. (2009). "Coat color variation at the beginning of horse domestication". Science. 324 (5926): 485. Bibcode:2009Sci...324..485L. doi:10.1126/science.1172750. PMC 5102060. PMID 19390039.
- Fang M.; et al. (2009). "Contrasting mode of evolution at a coat color locus in wild and domestic pigs". PLoS Genet. 5 (1): e1000341. doi:10.1371/journal.pgen.1000341. PMC 2613536. PMID 19148282.
- Hemmer, H. 1990. Domestication: The decline of environmental appreciation. Cambridge:Cambridge University Press
- Pennisi, E. (2015). "The taming of the pig took some wild turns". Science. doi:10.1126/science.aad1692.
- Li, Y. (2014). "Domestication of the dog from the wolf was promoted by enhanced excitatory synaptic plasticity: A hypothesis". Genome Biology and Evolution. 6 (11): 3115–21. doi:10.1093/gbe/evu245. PMC 4255776. PMID 25377939.
- Serpell J, Duffy D. Dog Breeds and Their Behavior. In: Domestic Dog Cognition and Behavior. Berlin, Heidelberg: Springer; 2014
- Cagan, Alex; Blass, Torsten (2016). "Identification of genomic variants putatively targeted by selection during dog domestication". BMC Evolutionary Biology. 16: 10. doi:10.1186/s12862-015-0579-7. PMC 4710014. PMID 26754411.
- Almada RC, Coimbra NC. Recruitment of striatonigral disinhibitory and nigrotectal inhibitory GABAergic pathways during the organization of defensive behavior by mice in a dangerous environment with the venomous snake Bothrops alternatus [ Reptilia, Viperidae ] Synapse 2015:n/a–n/a
- Coppinger R, Schneider R: Evolution of working dogs. The domestic dog: Its evolution, behaviour and interactions with people. Cambridge: Cambridge University press, 1995
- Pendleton, Amanda L.; Shen, Feichen; Taravella, Angela M.; Emery, Sarah; Veeramah, Krishna R.; Boyko, Adam R.; Kidd, Jeffrey M. (2018). "Comparison of village dog and wolf genomes highlights the role of the neural crest in dog domestication". BMC Biology. 16 (1): 64. doi:10.1186/s12915-018-0535-2. PMC 6022502. PMID 29950181.
- Arendt, M; Cairns, K M; Ballard, J W O; Savolainen, P; Axelsson, E (2016). "Diet adaptation in dog reflects spread of prehistoric agriculture". Heredity. 117 (5): 301–306. doi:10.1038/hdy.2016.48. PMC 5061917. PMID 27406651.
- Freedman, Adam H.; Schweizer, Rena M.; Ortega-Del Vecchyo, Diego; Han, Eunjung; Davis, Brian W.; Gronau, Ilan; Silva, Pedro M.; Galaverni, Marco; Fan, Zhenxin; Marx, Peter; Lorente-Galdos, Belen; Ramirez, Oscar; Hormozdiari, Farhad; Alkan, Can; Vilà, Carles; Squire, Kevin; Geffen, Eli; Kusak, Josip; Boyko, Adam R.; Parker, Heidi G.; Lee, Clarence; Tadigotla, Vasisht; Siepel, Adam; Bustamante, Carlos D.; Harkins, Timothy T.; Nelson, Stanley F.; Marques-Bonet, Tomas; Ostrander, Elaine A.; Wayne, Robert K.; Novembre, John (2016). "Demographically-Based Evaluation of Genomic Regions under Selection in Domestic Dogs". PLOS Genetics. 12 (3): e1005851. doi:10.1371/journal.pgen.1005851. PMC 4778760. PMID 26943675.
- Shipman 2015, pp. 149
- Shipman, Pat (2015). "How do you kill 86 mammoths? Taphonomic investigations of mammoth megasites". Quaternary International. 359–360: 38–46. Bibcode:2015QuInt.359...38S. doi:10.1016/j.quaint.2014.04.048.
- Zimov, S.A.; Zimov, N.S.; Tikhonov, A.N.; Chapin, F.S. (2012). "Mammoth steppe: A high-productivity phenomenon". Quaternary Science Reviews. 57: 26–45. Bibcode:2012QSRv...57...26Z. doi:10.1016/j.quascirev.2012.10.005.
- Hare, B. (2013). The Genius of Dogs. Penguin Publishing Group.
- Hare B. (2005). "Human-like social skills in dogs?". Trends in Cognitive Sciences. 9 (9): 439–44. doi:10.1016/j.tics.2005.07.003. PMID 16061417.
- Bruce Hood (psychologist) (2014). The Domesticated Brain. Pelican. ISBN 9780141974866.Preface
- Darcy, Morey (1994). "The Early Evolution of the Domestic Dog". American Scientist. 82 (4): 336–347. JSTOR 29775234.
- Perry, George H; Dominy, Nathaniel J; Claw, Katrina G; Lee, Arthur S; Fiegler, Heike; Redon, Richard; Werner, John; Villanea, Fernando A; Mountain, Joanna L; Misra, Rajeev; Carter, Nigel P; Lee, Charles; Stone, Anne C (2007). "Diet and the evolution of human amylase gene copy number variation". Nature Genetics. 39 (10): 1256–60. doi:10.1038/ng2123. PMC 2377015. PMID 17828263.
- Axelsson, Erik; Ratnakumar, Abhirami; Arendt, Maja-Louise; Maqbool, Khurram; Webster, Matthew T.; Perloski, Michele; Liberg, Olof; Arnemo, Jon M.; Hedhammar, Åke; Lindblad-Toh, Kerstin (2013). "The genomic signature of dog domestication reveals adaptation to a starch-rich diet". Nature. 495 (7441): 360–4. Bibcode:2013Natur.495..360A. doi:10.1038/nature11837. PMID 23354050.
- Butterworth, G. (2003). "Pointing is the royal road to language for babies".
- Lakatos, G. (2009). "A comparative approach to dogs' (Canis familiaris) and human infants' comprehension of various forms of pointing gestures". Animal Cognition. 12 (4): 621–31. doi:10.1007/s10071-009-0221-4. PMID 19343382.
- Muller, C. (2015). "Dogs can discriminate the emotional expressions of human faces". Current Biology. 25 (5): 601–5. doi:10.1016/j.cub.2014.12.055. PMID 25683806.
- Hare, B. (2013). "What Are Dogs Saying When They Bark?". Scientific American. Retrieved 17 March 2015.
- Sanderson, K. (2008). "Humans can judge a dog by its growl". Nature. doi:10.1038/news.2008.852.
- Nagasawa, M. (2015). "Oxytocin-gaze positive loop and the coevolution of human-dog bonds". Bibcode:2015Sci...348..333N. doi:10.1126/science.1261022.
- Paul Taçon, Pardoe, Colin (2002). "Dogs make us human". Nature Australia. Australian Museum. 27 (4): 52–61.CS1 maint: Uses authors parameter (link) also available: https://www.researchgate.net/publication/29464691_Dogs_make_us_human
- Temple Grandin and Catherine Johnson (2005). Animals in Translation: Using the Mysteries of Autism to Decode Animal Behavior (PDF). A Harvest Book, Harcourt, Inc. New York. p. 305.CS1 maint: Uses authors parameter (link)
- Kathryn Kirkpatrick (2014). Jeanne Dubino, Ziba Rashidian, Andrew Smyth (eds.). Representing the Modern Animal in Culture. Palgrave Macmillan New York. p. 75.CS1 maint: Uses editors parameter (link)
- Fogg, Brandy R.; Howe, Nimachia; Pierotti, Raymond (2015). "Relationships Between Indigenous American Peoples and Wolves 1: Wolves as Teachers and Guides". Journal of Ethnobiology. 35 (2): 262–285. doi:10.2993/etbi-35-02-262-285.1.
- Schleidt, W. M. (1998). "Is humaneness canine?". Human Ethology Bulletin. 13 (4): 1–4.
- Andrew Brown Smith (2005). African Herders: Emergence of Pastoral Traditions. Walnut Creek: Altamira Press. p. 27.
- Verworn, M., R. Bonnet, and G. Steinmann. 1914. Diluviale Menschenfunde in Obercassel bei Bonn. Naturwissenschaften 2 (27): 645–650. [Diluvial People found in Obercassel near Bonn]
- Verworn, M., R. Bonnet, G. Steinmann. 1919. Der diluviale Menschenfund von Obercassel bei Bonn. Wiesbaden. [The diluvial People found in Obercassel near Bonn]
- Nobis, G. 1979. Der älteste Haushund lebte vor 14 000 Jahren. Umschau 79 (19): 610.
- Nobis, G. 1981. Aus Bonn: Das älteste Haustier des Menschen. Unterkiefer eines Hundes aus dem Magdaleniengrab von Bonn-Oberkassel. Das Rheinische Landesmuseum Bonn: Berichte aus der Arbeit des Museums 4/81: 49–50.
- Benecke, Norbert (1987). "Studies on early dog remains from Northern Europe". Journal of Archaeological Science. 14: 31–49. doi:10.1016/S0305-4403(87)80004-3.
- Liane Giemsch, Susanne C. Feine, Kurt W. Alt, Qiaomei Fu, Corina Knipper, Johannes Krause, Sarah Lacy, Olaf Nehlich, Constanze Niess, Svante Pääbo, Alfred Pawlik, Michael P. Richards, Verena Schünemann, Martin Street, Olaf Thalmann, Johann Tinnes, Erik Trinkaus & Ralf W. Schmitz. "Interdisciplinary investigations of the late glacial double burial from Bonn-Oberkassel". Hugo Obermaier Society for Quaternary Research and Archaeology of the Stone Age: 57th Annual Meeting in Heidenheim, 7th – 11th April 2015, 36-37
- Janssens, Luc; Giemsch, Liane; Schmitz, Ralf; Street, Martin; Van Dongen, Stefan; Crombé, Philippe (2018). "A new look at an old dog: Bonn-Oberkassel reconsidered". Journal of Archaeological Science. 92: 126–138. doi:10.1016/j.jas.2018.01.004.
- Street, Martin & Janssens, Luc & Napierala, Hannes. (2015). Street, M., Napierala, H. & Janssens, L. 2015: The late Palaeolithic dog from Bonn-Oberkassel in context. In: The Late Glacial Burial from Oberkassel Revisited (L. Giemsch / R. W. Schmitz eds.), Rheinische Ausgrabungen 72, 253-274. ISBN 978-3-8053-4970-3. Rheinische Ausgrabungen.
- Coppinger R, Feinstein M (2015). How Dogs Work. University of Chicago Press, Chicago. ISBN 9780226128139.CS1 maint: Uses authors parameter (link)
- Davis, S (1982). "The taming of the few". New Scientist. 95: 697–700.
- Clutton-Brock, J. 1984. Dog, in I.L. Mason (ed.) Evolution of domesticated animals. London:Longman.
- Perri, Angela R. (2016). "Hunting dogs as environmental adaptations in Jōmon Japan". Antiquity. 90 (353): 1166–1180. doi:10.15184/aqy.2016.115.
- Guagnin, Maria; Perri, Angela R.; Petraglia, Michael D. (2018). "Pre-Neolithic evidence for dog-assisted hunting strategies in Arabia". Journal of Anthropological Archaeology. 49: 225–236. doi:10.1016/j.jaa.2017.10.003.
- Perri, Angela; Widga, Chris; Lawler, Dennis; Martin, Terrance; Loebel, Thomas; Farnsworth, Kenneth; Kohn, Luci; Buenger, Brent (2019). "New Evidence of the Earliest Domestic Dogs in the Americas". American Antiquity. 84: 68–87. doi:10.1017/aaq.2018.74.
- Ní Leathlobhair, Máire; Perri, Angela R; Irving-Pease, Evan K; Witt, Kelsey E; Linderholm, Anna; Haile, James; Lebrasseur, Ophelie; Ameen, Carly; Blick, Jeffrey; Boyko, Adam R; Brace, Selina; Cortes, Yahaira Nunes; Crockford, Susan J; Devault, Alison; Dimopoulos, Evangelos A; Eldridge, Morley; Enk, Jacob; Gopalakrishnan, Shyam; Gori, Kevin; Grimes, Vaughan; Guiry, Eric; Hansen, Anders J; Hulme-Beaman, Ardern; Johnson, John; Kitchen, Andrew; Kasparov, Aleksei K; Kwon, Young-Mi; Nikolskiy, Pavel A; Lope, Carlos Peraza; et al. (2018). "The evolutionary history of dogs in the Americas". Science. 361 (6397): 81–85. doi:10.1126/science.aao4776. PMID 29976825.
- Pitulko, Vladimir V.; Kasparov, Aleksey K. (2017). "Archaeological dogs from the Early Holocene Zhokhov site in the Eastern Siberian Arctic". Journal of Archaeological Science: Reports. 13: 491–515. doi:10.1016/j.jasrep.2017.04.003.
- Ikeya, K. 1994. Hunting with dogs among the San in the Central Kalahari. African Study Monographs 15:119–34.
- Gron, O. & M.G. Turov. 2007. Resource 'pooling' and resource management. An ethno-archaeological study of the Evenk hunter-gatherers, Katanga County, Siberia, in B. Hardh, K. Jennbert & D. Olausson (ed.) On the road: studies in honour of Lars Larsson (Acta Archaeologica Lundensia 26):67–72. Stockholm: Almqvist & Wiksell.
- Koler-Matznick, Janice; Brisbin Jr, I. Lehr; Yates, S; Bulmer, Susan (2007). "The New Guinea singing dog: its status and scientific importance". The Journal of the Australian Mammal Society. 29 (1): 47–56. CiteSeerX 10.1.1.627.5761. doi:10.1071/AM07005.
- Olowo Ojoade, J. 1990. Nigerian cultural attitudes to the dog, in R. Willis (ed.) Signifying animals: human meaning in the natural world: 215–21. London: Routledge.
- Mizoguchi, K. 2002. An archaeological history of Japan: 10,000 B.C. to A.D. 700. Philadelphia: University of Pennsylvania Press.
- Bourque, B.J. 1975. Comments on the late Archaic populations of central Maine: the view from the urner Farm. Arctic Anthropology 12: 35–45.
- Larsson, L. 1990. Dogs in fraction—symbols in action, in P.M. Vermeersch & P. Van Peer (ed.) Contributions to the Mesolithic in Europe: 153–60. Leuven: Leuven University Press.
- Morey, D.F. (1992). "Early Holocene domestic dog burials from the North American Midwest". Current Anthropology. 33 (2): 224–29. doi:10.1086/204059.
- Derr, Mark (2011). How the Dog Became the Dog: From Wolves to Our Best Friends. Penguin Group. ISBN 978-1468302691.
- Pierotti, R.; Fogg, B. (2017). The First Domestication: How Wolves and Humans Coevolved. Yale University Press. ISBN 978-0-300-22616-4.
- Shipman, P. (2015). The Invaders:How humans and their dogs drove Neanderthals to extinction. Harvard University Press. ISBN 9780674736764.