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The grey wolf (Canis lupus) was the first species to give rise to a domestic population, and they remained widespread throughout the last Ice Age when many other large mammal species went extinct. Little is known, however, about the history and possible extinction of past wolf populations or when and where the wolf progenitors of the present-day dog lineage (Canis familiaris) lived1–8. Here we analysed 72 ancient wolf genomes spanning the last 100,000 years from Europe, Siberia and North America. We found that wolf populations were highly connected throughout the Late Pleistocene, with levels of differentiation an order of magnitude lower than they are today. This population connectivity allowed us to detect natural selection across the time series, including rapid fixation of mutations in the gene IFT88 40,000–30,000 years ago. We show that dogs are overall more closely related to ancient wolves from eastern Eurasia than to those from western Eurasia, suggesting a domestication process in the east. However, we also found that dogs in the Near East and Africa derive up to half of their ancestry from a distinct population related to modern southwest Eurasian wolves, reflecting either an independent domestication process or admixture from local wolves. None of the analysed ancient wolf genomes is a direct match for either of these dog ancestries, meaning that the exact progenitor populations remain to be located.

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The Bonn-Oberkassel dog remains (Upper Pleistocene and 14223 þ-58 years old) have been reported more than 100 years ago. Recent re-examination revealed the tooth of another older and smaller dog, making this domestic dog burial not only the oldest known, but also the only one with remains of two dogs. This observation brings the total known Magdalenian dogs to nine. Domestication of dogs during the final Palaeolithic has important implications for understanding pre-Holocene hunter-gatherers. Most proposed hunter-gatherer motivations for domesticating dogs have been utilitarian. However, remains of the Bonn-Oberkassel dogs may offer another view. The Bonn-Oberkassel dog was a late juvenile when it was buried at approximately age 27e28 weeks, with two adult humans and grave goods. Oral cavity lesions indicate a gravely ill dog that likely suffered a morbillivirus (canine distemper) infection. A dental line of suggestive enamel hypoplasia appears at the 19-week developmental stage. Two additional enamel hypoplasia lines, on the canine only, document further disease episodes at weeks 21 and 23. Pathological changes also include severe periodontal disease that may have been facilitated by immunodeficiency. Since canine distemper has a three-week disease course with very high mortality, the dog must have been perniciously ill during the three disease bouts and between ages 19 and 23 weeks. Survival without intensive human assistance would have been unlikely. Before and during this period, the dog cannot have held any utilitarian use to humans. We suggest that at least some Late Pleistocene humans regarded dogs not just materialistically, but may have developed emotional and caring bonds for their dogs, as reflected by the survival of this dog, quite possibly through human care.

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