Tracking the evolutionary origins of dog-human cooperation: the “Canine Cooperation Hypothesis”
According to Groves: "The human-dog relationship amounts to a very long lasting symbiosis. Dogs Evolution and Human Behavior, 18, 2. Udell. Article: History & Evolution» Dogs and Humans  The relationship between the presence of a dog and success in the hunt is often mentioned as a primary. Keywords: domestication, evolution, cooperation, attention, Canis lupus, Dogs' attention towards humans depends on their relationship, not.
The sample provided the first draft genome from the cell nucleus of a Pleistocene carnivore and the sequence was identified as belonging to Canis lupus. The sequence indicated that the Taimyr-1 lineage was separate to modern wolves and dogs. Using the Taimyr-1 specimen's radiocarbon date in addition to its genome sequence compared to that of a modern wolf, a direct estimate of the mutation rate in dogs and wolves could be made to calculate the time of divergence.The Origin of Dogs
The study calculated a mutation rate for the 7, YBP Neolithic dog and found that it matched the mutation rate of the Taimyr-1 specimen, and noted that this also matched the mutation rate for the Newgrange dog that had been calculated in an earlier study. Using the 7, YBP specimen and this mutation rate, the dog-wolf divergence time is estimated to have occurred 36, YBP and this is consistent with the timing found with the Taimyr-1 specimen in an earlier study.
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, YBP.
Studies of modern grey wolves have identified distinct sub-populations that live in close proximity to each other.
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Where the genetic divergence of dog and wolf took place remains controversial, with the most plausible proposals spanning Western Europe,   Central Asia,   and East Asia. Ina study of the maternal mitochondrial genome indicated the origin in south-eastern Asia south of the Yangtze River as more dog haplogroups had been found there. Ina study using single nucleotide polymorphisms indicated that dogs originated in the Middle East due to the greater sharing of haplotypes between dogs and Middle Eastern gray wolves, else there may have been significant admixture between some regional breeds and regional wolves.
Ina study of maternal mDNA indicated that the dog diverged from its ancestor in East Asia because there were more dog mDNA haplotypes found there than in other parts of the world,  but this was rebutted because village dogs in Africa also show a similar haplotype diversity.
Then, one of these lineages migrated back to northern China and admixed with endemic Asian lineages before migrating to the Americas. Ina study looked at 85, genetic markers of autosomalmaternal mitochondrial genome and paternal Y chromosome diversity in 4, purebred dogs from breeds and village dogs from 38 countries. Some dog populations in the Neotropics and the South Pacific are almost completely derived from European stock, and other regions show clear admixture between indigenous and European dogs.
The indigenous dog populations of Vietnam, India, and Egypt show minimal evidence of European admixture, and exhibit indicators consistent with a Central Asian domestication origin, followed by a population expansion in East Asia.
Tracking the evolutionary origins of dog-human cooperation: the “Canine Cooperation Hypothesis”
The study could not rule out the possibility that dogs were domesticated elsewhere and subsequently arrived in and diversified from Central Asia. Studies of extant dogs cannot exclude the possibility of earlier domestication events that subsequently died out or were overwhelmed by more modern populations. Ina whole-genome study of wolves and dogs concluded that admixture had confounded the ability to make inferences about the place of dog domestication.
Past studies based on single-nucleotide polymorphisms genome-wide similarities with Chinese wolves,  and lower linkage disequilibrium  might reflect regional admixture between dogs with wolves and gene flow between dog populations, with genetically divergent dog breeds possibly maintaining more wolf ancestry in their genome. The study proposed that the analysis of ancient DNA might be a better approach.
The advent of rapid and inexpensive DNA sequencing technology has made it possible to significantly increase the resolving power of genetic data taken from both modern and ancient domestic dog genomes. Attention was now turned to studies based on ancient DNA from fossil canids. Ina study analysed the complete and partial mitochondrial genome sequences of 18 fossil canids dated from 1, to 36, YBP from the Old and New Worlds, and compared these with the complete mitochondrial genome sequences from modern wolves and dogs.
Phylogenetic analysis showed 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. This group of dogs matched three fossil pre-Columbian New World dogs dated between 1, and 8, YBP, which supported the hypothesis that pre-Columbian dogs in the New World share ancestry with modern dogs and that they likely arrived with the first humans to the New World.
However, this relationship might represent mitochondrial genome introgression from wolves because dogs were domesticated by this time.
Clade D contained sequences from 2 Scandinavian breeds JamthundNorwegian Elkhound and were sister to another 14, YBP wolf sequence also from the Kesserloch cave, with a common recent ancestor estimated to 18, YBP. Its branch is phylogenetically rooted in the same sequence as the "Altai dog" not a direct ancestor. The study found that the skulls of the "Goyet dog" and the "Altai dog" had some dog-like characteristics and proposed that the may have represented an aborted domestication episode.
If so, there may have been originally more than one ancient domestication event for dogs  as there was for domestic pigs.
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The theory is that the extreme cold during one of these events caused humans to either shift their location, adapt through a breakdown in their culture and change of their beliefs, or adopt innovative approaches.
However, dramatic differences in genetic diversity can be influenced both by an ancient and recent history of inbreeding. Ina study looked at the mitochondrial control region sequences of 13 ancient canid remains and one modern wolf from five sites across Arctic north-east Siberia. The fourteen canids revealed nine mitochondrial haplotypesthree of which were on record and the others not reported before.
This article was submitted to the journal Frontiers in Psychology. Received Jul 24; Accepted Dec The use, distribution or reproduction in other forums is permitted, provided the original author s or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
This article has been cited by other articles in PMC. Abstract At present, beyond the fact that dogs can be easier socialized with humans than wolves, we know little about the motivational and cognitive effects of domestication. Despite this, it has been suggested that during domestication dogs have become socially more tolerant and attentive than wolves.
These two characteristics are crucial for cooperation, and it has been argued that these changes allowed dogs to successfully live and work with humans. However, these domestication hypotheses have been put forward mainly based on dog-wolf differences reported in regard to their interactions with humans.
Thus, it is possible that these differences reflect only an improved capability of dogs to accept humans as social partners instead of an increase of their general tolerance, attentiveness and cooperativeness.
At the Wolf Science Center, in order to detangle these two explanations, we raise and keep dogs and wolves similarly socializing them with conspecifics and humans and then test them in interactions not just with humans but also conspecifics. When investigating attentiveness toward human and conspecific partners using different paradigms, we found that the wolves were at least as attentive as the dogs to their social partners and their actions.
Based on these findings and the social ecology of wolves, we propose the Canine Cooperation Hypothesis suggesting that wolves are characterized with high social attentiveness and tolerance and are highly cooperative. This is in contrast with the implications of most domestication hypotheses about wolves. We argue, however, that these characteristics of wolves likely provided a good basis for the evolution of dog-human cooperation.
Although it is clear that human collaborative skills are exceptional, studying the cognitive and emotional processes of animal species that may underlie their cooperative interactions may reveal the evolutionary origins and the functional relevance of cooperation. For a long time, the most common approach to investigate the evolutionary origin of human skills was to study non-human primates.
This idea is built on the assumption that dogs have been selected to cooperate and communicate with humans during domestication and, thus, evolved some genetic predispositions allowing them to develop skills shared with humans Hare et al.
For instance, it has been argued that bonobos Pan paniscus outperform chimpanzees Pan troglodytes in cooperative interactions Hare et al. Moreover, also at an individual level, tolerant individuals usually outperform less tolerant ones in cooperative tasks marmosets Callithrix jacchus: Werdenich and Huber, ; chimpanzees: Chalmeau and Gallo, ; Melis et al.
Along the lines of this argument, Hare and Tomasello proposed that selection for a tamer temperament and for reduced fear and aggression explains the higher success of dogs in cooperative and communicative interactions with humans in comparison to wolves, the closest wild-living relative of dogs emotional reactivity hypothesis.
Recently, this hypothesis has been extended to suggest that during domestication dogs became less aggressive and more tolerant than wolves not just toward humans but also toward conspecifics Hare et al.