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Published July 7, 2021 | Version 1.0.1
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The Origins of SARS-CoV-2: A Critical Review

  • 1. School of Life and Environmental Sciences and School of Medical Sciences, The University of Sydney, Sydney, NSW 2006, Australia
  • 2. Department of Human Genetics, University of Utah, Salt Lake City, UT 84112, USA
  • 3. Vaccine and Infectious Disease Organization, University of Saskatchewan, Saskatoon, SK, S7N 5E3, Canada
  • 4. MRC-University of Glasgow Centre for Virus Research, Glasgow, G61 1QH, UK
  • 5. Department of Plant and Microbial Biology, University of California Berkeley, Berkeley, CA 94704, USA
  • 6. Department of Medicine, University of California San Diego, La Jolla, CA 92093, USA
  • 7. Department of Pathology, Microbiology, and Immunology, University of California Davis School of Veterinary Medicine, Davis, CA 95616, USA
  • 8. Department of Infectious Disease, Imperial College London, W2 1PG, UK
  • 9. Center for Infectious Disease Dynamics, Department of Biology, The Pennsylvania State University, University Park, PA 16802, USA
  • 10. Department of Microbiology and Immunology, The University of Melbourne at the Doherty Institute, 792 Elizabeth St, Melbourne, VIC 3000, Australia
  • 11. The Wellcome Trust, London, NW1 2BE, UK
  • 12. Department of Microbiology and Immunology, University of Otago, Dunedin, New Zealand. Institute of Environmental Science and Research, Wellington, New Zealand
  • 13. Department of Biological Sciences, Xi'an Jiaotong-Liverpool University (XJTLU), Suzhou, China
  • 14. Department of Microbial Pathogenesis and Immunology, Texas A&M University, College Station, TX 77807, USA
  • 15. Department of Infectious Diseases, King's College London, Guy's Hospital, London SE1 9RT, UK
  • 16. Max Perutz Labs, Medical University of Vienna, Vienna Biocenter, Dr. Bohr-Gasse 9/3, A-1030 Vienna, Austria
  • 17. Perelman School of Medicine, University of Pennsylvania. Philadelphia, PA 19104, USA
  • 18. Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, USA
  • 19. Department of Immunology and Microbiology, The Scripps Research Institute, La Jolla, CA 92037, USA
  • 20. Department of Microbiology and Immunology, Tulane University School of Medicine, New Orleans, LA 70112, USA
  • 21. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh, EH9 FL, UK


The Origins of SARS-CoV-2: A Critical Review

Holmes et al.

Since the first reports of a novel SARS-like coronavirus in December 2019 in Wuhan, China, there has been intense interest in understanding how SARS-CoV-2 emerged in the human population. Recent debate has coalesced around two competing ideas: a “laboratory escape” scenario and zoonotic emergence. Here, we critically review the current scientific evidence that may help clarify the origin of SARS-CoV-2.

Computer readable versions of data tables, SVG maps, and acknowledgements of sequence data used are available from:



v1.0.1 includes some adjustments to the maps so the points better capture the maps in the WHO report annex.



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