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Dire wolves were the last of an ancient New World canid lineage


Dire wolves are considered to be one of the most common and widespread large carnivores in Pleistocene America1, yet relatively little is known about their evolution or extinction. Here, to reconstruct the evolutionary history of dire wolves, we sequenced five genomes from sub-fossil remains dating from 13,000 to more than 50,000 years ago. Our results indicate that although they were similar morphologically to the extant grey wolf, dire wolves were a highly divergent lineage that split from living canids around 5.7 million years ago. In contrast to numerous examples of hybridization across Canidae2,3, there is no evidence for gene flow between dire wolves and either North American grey wolves or coyotes. This suggests that dire wolves evolved in isolation from the Pleistocene ancestors of these species. Our results also support an early New World origin of dire wolves, while the ancestors of grey wolves, coyotes and dholes evolved in Eurasia and colonized North America only relatively recently.

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Fig. 1: Map of dire wolf remains and morphological differentiation with wolf-like canids.
Fig. 2: Relationships among living and extinct wolf-like canids.

Data availability

The reads generated for this study have been deposited in the European Nucleotide Archive (ENA) (project number PRJEB31639). The accession numbers for the publicly available genomes used in this study can be found in Supplementary Table 2 and Supplementary Data 13. The mass-spectrometry proteomics data have been deposited in the ProteomeXchange Consortium via the PRIDE partner repository (PXD021930). Ancient collagen consensus sequences for the dire wolf can be found in Supplementary Data 17. Two-dimensional mandibular and dental shape (geometric morphometric) data have been deposited in Dryad (


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We thank the staff at the Carnegie Museum of Natural History, Cincinnati Museum Center, Danish Zoological Museum, Harrison Zoological Museum, Harvard Museum of Comparative Zoology, Idaho Museum of Natural History, Institute of Archaeology (Russian Academy of Sciences), Institute of Systematics and Animal Ecology (Russian Academy of Sciences), Institute of Zoology (Chinese Academy of Sciences), Instituto de Conservação da Natureza e das Florestas, Kansas Museum of Natural History, La Brea Tar Pits and Museum, Ludwig Maximilian University, McClung Museum, Museum of the Institute of Plant and Animal Ecology (Russian Academy of Sciences), Museum national d’Histoire naturelle, National Museums Scotland, Natural History Museum London, Naturalis Biodiversity Center, Naturhistorisches Museum Bern, Smithsonian National Museum of Natural History, Swedish Naturhistoriska Riksmuseet, SYLVATROP, US Bureau of Reclamation, University of California Museum of Paleontology, University of Texas at El Paso, University of Washington Burke Museum and the Zoological Institute (Russian Academy of Sciences; state assignment no. АААА-А19-119032590102-7) for access to specimens in their care; T. Barnosky, S. Bray, A. Farrell, R. Fischer, A. Harris, J. Harris, A. Henrici, P. Holroyd, R. MacPhee, T. Martin, A. Philpot, J. Saunders, J. Southon, G. Storrs, G. Takeuchi, X. Wang and C. Widga for assistance; and L. DeSantis for comments. A.M. used computational and storage services associated with the Hoffman2 Shared Cluster provided by UCLA Institute for Digital Research and Education’s Research Technology Group. DireGWC was sequenced using the Vincent J. Coates Genomics Sequencing Laboratory at UC Berkeley, supported by NIH S10 OD018174 Instrumentation Grant. We acknowledge the assistance of the Danish National High-Throughput Sequencing Centre, BGI-Europe, the Garvan Institute of Medical Research and the Australian Cancer Research Foundation (ACRF) Cancer Genomics Facility for assistance in Illumina and BGIseq500 data generation. A.R.P. was supported by a Marie Curie COFUND Junior Research Fellowship (Durham University). A.M. was supported by an NSF grant (award number: 1457106) and the QCB Collaboratory Postdoctoral Fellowship (UCLA). L.A.F.F., J.H., A.H.-B. and G.L. were supported by either European Research Council grant (ERC-2013-StG-337574-UNDEAD and ERC-2019-StG-853272-PALAEOFARM) and/or Natural Environmental Research Council grants (NE/K005243/1 and NE/K003259/1). K.S. was supported by a grant from Barrett, the Honors College at Arizona State University. A.T.O. was supported by the Strategic Initiative Funds, Office of the President, Arizona State University to the Institute of Human Origins DNA and Human Origins at Arizona State University project. L.A.F.F. was supported by a Junior Research Fellowship (Wolfson College, University of Oxford) and L.A.F.F. and A. Carmagnini were supported by the Wellcome Trust (210119/Z/18/Z). S.G. was supported by Carlsbergfondet grant CF14–0995 and Marie Skłodowska-Curie Actions grant 655732-WhereWolf. M.T.P.G. was supported by ERC Consolidator grant 681396-Extinction Genomics. B.S. and J.K. were supported by IMLS MG-30-17-0045-17 and NSF DEB-1754451. A.H.-B. was supported by the Leverhulme Trust (ECF-2017-315). A. Cooper, K.J.M. and H.H. were supported by the Australian Research Council. A.T.S. and G.G. were supported by Australian Government Research Training Program Scholarships. A.T.L. was supported by the Peter Buck Postdoctoral Fellowship from the Smithsonian Institution’s National Museum of Natural History. Y.V.K. was supported by the by State Assignment of the Sobolev Institute of Geology and Mineralogy.

Author information




A.R.P., K.J.M., A.M., R.K.W., G.L., L.A.F.F. and A. Cooper conceived the project and designed the research; A.R.P. and K.J.M. coordinated the sample collection efforts with input from R.K.W., G.L., L.A.F.F. and A. Cooper; A.R.P., K.J.M., A.H.-B., J.M., C.A., J.C.C., A.E., P.G., J.K., A.L., A.T.O., S.P., B.W.S., M.T., M.J.C., M.-H.S.S., M.T.P.G., A.C.S., B.S., B.V.V., R.W.K. and A. Cooper provided and/or collected samples; A.R.P., K.J.M., R.K.W., A.M., C.S., J.H., A.J., A.T.S., P.B. and H.H. conducted the genomic laboratory work; K.J.M., A.M., G.G., G.L., L.A.F.F. and A. Cooper conducted the analyses of the genomic data; S.A.-C., A.H.-B., J.M., C.A., K.M.H., and A.E. conducted the morphological analyses; A.R.P., K.J.M., A.M., S.A.-C., B.V.V., K.M.H., R.K.W., G.L., L.A.F.F. and A. Cooper wrote the paper with input from all other authors.

Corresponding authors

Correspondence to Angela R. Perri or Kieren J. Mitchell or Laurent A. F. Frantz.

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The authors declare no competing interests.

Additional information

Peer review information Nature thanks Larisa DeSantis and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Peer reviewer reports are available.

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Supplementary information

Supplementary Information

This file contains Supplementary Methods, Supplementary Discussion, Supplementary Figures 1-23, and Supplementary Tables 1-11.

Reporting Summary

Peer Review File

Supplementary Data

This file contains Supplementary Data 1-16.

Supplementary Data

This file contains the collagen amino sequence of the dire wolf in a fasta format.

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Perri, A.R., Mitchell, K.J., Mouton, A. et al. Dire wolves were the last of an ancient New World canid lineage. Nature 591, 87–91 (2021).

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