Ancient Wolf Genome Reveals an Early Divergence of Domestic Dog Ancestors and Admixture into High-Latitude Breeds  Pontus Skoglund, Erik Ersmark, Eleftheria.

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Ancient Wolf Genome Reveals an Early Divergence of Domestic Dog Ancestors and Admixture into High-Latitude Breeds  Pontus Skoglund, Erik Ersmark, Eleftheria Palkopoulou, Love Dalén  Current Biology  Volume 25, Issue 11, Pages 1515-1519 (June 2015) DOI: 10.1016/j.cub.2015.04.019 Copyright © 2015 Elsevier Ltd Terms and Conditions

Figure 1 Mitochondrial DNA Phylogeny The outgroups (one coyote and two Chinese wolf sequences) were excluded. Nodes with posterior probability >0.9 are marked with asterisks, and stars indicate ancient samples morphologically described as “dog like.” See also Figure S1 and Table S1. Current Biology 2015 25, 1515-1519DOI: (10.1016/j.cub.2015.04.019) Copyright © 2015 Elsevier Ltd Terms and Conditions

Figure 2 The 35,000 Year-Old Taimyr Individual Belonged to a Population that Was Genetically Close to the Ancestor of Present-Day Gray Wolves and Domestic Dogs (A) D statistics testing the phylogenetic relationship between the Taimyr wolf and present-day canids. The Andean fox is used as an outgroup in all tests. Error bars correspond to one block jackknife SE on each side. (B) A model of population history (admixture graph) fitted to the data. The Andean fox, which was used as an outgroup, is not shown. Although the graph shown here indicates minor gene flow from domestic dogs into regional wolf populations, we note that an alternative graph with gene flow from the wolf lineages into the domestic dog lineages provides equally good fit. In all fitted graphs, the branch length between the divergence of the Taimyr wolf lineage and the wolf-dog split is very short. Branch lengths are measured as the allele frequency variance along each branch (genetic drift) rescaled to be proportional to FST. See also Figures S2 and S3. Current Biology 2015 25, 1515-1519DOI: (10.1016/j.cub.2015.04.019) Copyright © 2015 Elsevier Ltd Terms and Conditions

Figure 3 Recalibrating the Lupine Mutation Rate using the Directly Dated Taimyr Genome We estimated the probability F(A|B) for the Taimyr individual carrying the derived allele at positions where the Chinese wolf is heterozygous using empirical data. This statistic is informative on the coalescent time passed in the ancestry of the Chinese wolf since its divergence from the Taimyr wolf’s lineage. Since the Taimyr wolf must have separated at least 35,000 years ago, the age of the specimen obtained from a direct radiocarbon date, its divergence from the Chinese wolf can be used to calibrate the lupine mutation rate, in the sense that we can infer the maximum mutation rate that is consistent with the proximity of the Taimyr genome to the present-day Chinese wolf. To achieve this, we built calibration curves for F(A|B) given a model of Chinese wolf effective population size history (inferred using pairwise sequentially Markovian coalescent [PSMC] analysis) and different mutation rates. Mutation rates per generation are shown in the legend. A generation time of 3 years is assumed. See also Table S2. Current Biology 2015 25, 1515-1519DOI: (10.1016/j.cub.2015.04.019) Copyright © 2015 Elsevier Ltd Terms and Conditions

Figure 4 Introgression from a Population Related to the Ancient Taimyr Wolf into Northeast Siberian and New World Arctic Dog Breeds We computed the statistic D(ancient Taimyr wolf, present-day gray wolves; dog breed X, pool of European breeds) where the pool of European breeds was comprised of 83 Boxer, Gordon Setter, Doberman Pinscher, and Newfoundland individuals. The color scale represents Z scores where the obtained D statistic values were normalized by SEs estimated using a block jackknife. Positive statistics suggest an excess affinity to the Taimyr wolf over the modern-day gray wolves for dog breed X. Negative statistics suggest an excess affinity to present-day gray wolves over the ancient Taimyr wolf for dog breed X. We tested 48 different dog breeds with information from ∼66,000 SNPs. The affinity of the Siberian Husky and Greenland Sledge Dogs to Taimyr is replicated also in statistics of the form D(Andean fox, Taimyr; present-day gray wolves, dog breed X). See also Figure S4 and Tables S3 and S4. Current Biology 2015 25, 1515-1519DOI: (10.1016/j.cub.2015.04.019) Copyright © 2015 Elsevier Ltd Terms and Conditions

Current Biology 2015 25, 1515-1519DOI: (10.1016/j.cub.2015.04.019) Copyright © 2015 Elsevier Ltd Terms and Conditions