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Evolutionary origins of the SARS-CoV-2 sarbecovirus lineage responsible for the COVID-19 pandemic
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Abstract
There are outstanding evolutionary questions on the recent emergence of human coronavirus SARS-CoV-2 including the role of reservoir species, the role of recombination and its time of divergence from animal viruses. We find that the sarbecoviruses-the viral subgenus containing SARS-CoV and SARS-CoV-2-undergo frequent recombination and exhibit spatially structured genetic diversity on a regional scale in China. SARS-CoV-2 itself is not a recombinant of any sarbecoviruses detected to date, and its receptor-binding motif, important for specificity to human ACE2 receptors, appears to be an ancestral trait shared with bat viruses and not one acquired recently via recombination. To employ phylogenetic dating methods, recombinant regions of a 68-genome sarbecovirus alignment were removed with three independent methods. Bayesian evolutionary rate and divergence date estimates were shown to be consistent for these three approaches and for two different prior specifications of evolutionary rates based on HCoV-OC43 and MERS-CoV. Divergence dates between SARS-CoV-2 and the bat sarbecovirus reservoir were estimated as 1948 (95% highest posterior density (HPD): 1879-1999), 1969 (95% HPD: 1930-2000) and 1982 (95% HPD: 1948-2009), indicating that the lineage giving rise to SARS-CoV-2 has been circulating unnoticed in bats for decades.
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“Although the proximal origins are unknown, SARS-CoV-2 is thought to derive from a bat SARS-CoV-related coronavirus with a furin cleavage site that enhances the capacity of the virus to infect human cells.48, 49”
From The Lancet Commission on lessons for the future from the COVID-19 pandemic (2022) · cites this paper as [48]
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The COVID-19 pandemic is intricately linked to biodiversity loss and ecosystem health
Lawler, O. K., Allan, H., Baxter, P. W. J., Castagnino, R., Tor, M. C., Dann, L. E., Hungerford, J., Karmacharya, D. B., Lloyd, T. J., López, M. J., et al. (2021) · The Lancet Planetary Health · doi:10.1016/s2542-5196(21)00258-8 · Cited by 204
“14,16 Direct spillover of SARS-CoV-2 or its progenitor to humans from a bat host has also been proposed as a possibility (appendix pp 1–2).”
BackgroundThe Evolution and Biology of SARS-CoV-2 VariantsPreprint
Telenti, A., Hodcroft, E. B., & Robertson, D. L. (2022) · Cold Spring Harbor Perspectives in Medicine · doi:10.1101/cshperspect.a041390 · Cited by 191
Self-citationTriggle, C. R., Bansal, D., Ding, H., Islam, M. M., Farag, E., Hadi, H. A., & Sultan, A. A. (2021) · Frontiers in Immunology · doi:10.3389/fimmu.2021.631139 · Cited by 187
“Genetic sequence data reveals that SARS-CoV-2 is a close relative of other species of coronaviruses found to be circulating in Rhinolophus bat (Horseshoe Bat) populations (9, 16).”
BackgroundExploring the Natural Origins of SARS-CoV-2 in the Light of Recombination
Lytras, S., Hughes, J., Martin, D. P., Swanepoel, P., de Klerk, A., Lourens, R., Pond, S. L. K., Xia, W., Jiang, X., & Robertson, D. L. (2022) · Genome Biology and Evolution · doi:10.1093/gbe/evac018 · Cited by 174
Self-citationAnimal and translational models of SARS-CoV-2 infection and COVID-19
Johansen, M. D., Irving, A. T., Montagutelli, X., Tate, M. D., Rudloff, I., Nold, M. F., Hansbro, N. G., Kim, R., Donovan, C., Liu, G., et al. (2020) · Mucosal Immunology · doi:10.1038/s41385-020-00340-z · Cited by 173
“OTHER INFECTED ANIMALS Bats While SARS-CoV-2 is clearly a suitable human virus, its origins remains elusive with a common ancestor of bat SARS-Like CoVs and SARS-CoV-2 existing ~40–70 years prior to emergence.(155) Scientists are now searching for further evidence in wild bats and have performed infection studies with…”
BackgroundPolymeric nanoparticle vaccines to combat emerging and pandemic threats
Wibowo, D., Jorritsma, S. H. T., Gonzaga, Z. J., Evert, B. J., Chen, S., & Rehm, B. H. A. (2020) · Biomaterials · doi:10.1016/j.biomaterials.2020.120597 · Cited by 177
“A notable example is the ongoing pandemic coronavirus disease 2019 (COVID-19) caused by a novel severe acute respiratory syndrome coronavirus-2 (SARS-CoV-2) originated from bats [4].”
Timing the SARS-CoV-2 index case in Hubei Province
Pekar, J., Worobey, M., Moshiri, N., Scheffler, K., & Wertheim, J. O. (2021) · Science · doi:10.1126/science.abf8003 · Cited by 156
Olival, K. J., Cryan, P. M., Amman, B. R., Baric, R. S., Blehert, D. S., Brook, C. E., Calisher, C. H., Castle, K. T., Coleman, J. T. H., Daszak, P., et al. (2020) · PLoS Pathogens · doi:10.1371/journal.ppat.1008758 · Cited by 172
“Third, if SARS-CoV-2 infection persists in North American bats of one or more species, it could potentially evolve or recombine with endemic viruses [19, 21] to become more pathogenic or infectious to humans or other animals.”
BackgroundHigh-throughput sequencing of SARS-CoV-2 in wastewater provides insights into circulating variants
Fontenele, R. S., Kraberger, S. J., Hadfield, J., Driver, E. M., Bowes, D. A., Holland, L. A., Faleye, T. O. C., Adhikari, S., Kumar, R., Inchausti, R., et al. (2021) · Water Research · doi:10.1016/j.watres.2021.117710 · Cited by 167
Self-citationControl and prevention of infectious diseases from a One Health perspective
Ellwanger, J. H., da Veiga, A. B. G., Kaminski, V. D. L., Valverde-Villegas, J. M., de Freitas, A. W. Q., & Chies, J. A. B. (2021) · Genetics and Molecular Biology · doi:10.1590/1678-4685-gmb-2020-0256 · Cited by 200
Profile of a killer: the complex biology powering the coronavirus pandemic
Cyranoski, D. (2020) · Nature · doi:10.1038/d41586-020-01315-7 · Cited by 170
An engineered decoy receptor for SARS-CoV-2 broadly binds protein S sequence variants
Chan, K. K., Tan, T. J. C., Narayanan, K. K., & Procko, E. (2021) · Science Advances · doi:10.1126/sciadv.abf1738 · Cited by 152
“In addition, large changes in coronavirus genomes have frequently occurred in nature from recombination events, especially in bats where coinfection levels can be high (15, 16).”
BackgroundNchioua, R., Kmiec, D., Müller, J. A., Conzelmann, C., Groß, R., Swanson, C. M., Neil, S. J. D., Stenger, S., Sauter, D., Münch, J., et al. (2020) · mBio · doi:10.1128/mbio.01930-20 · Cited by 146
“In this part of the genome, SARS-CoV-2 shows substantially higher similarity in sequence and CpG numbers to bat RaTG13 and pangolin Pang Guangdong CoV strains than to RmYN02 (20, 54).”
BackgroundInfluentialTransmission of SARS-CoV-2 from humans to animals and potential host adaptation
Tan, C. C. S., Lam, S. D., Richard, D., Owen, C. J., Berchtold, D., Orengo, C. A., Nair, M. S., Kuchipudi, S. V., Kapur, V., van Dorp, L., et al. (2022) · Nature Communications · doi:10.1038/s41467-022-30698-6 · Cited by 154
“Evolutionary analyses of SARS-CoV-2 and close relatives suggest that both ef fi cient human-to-human transmission and ACE2 usage were not acquired recently, but may already have been present in ancestral bat-associated lineages 4,5 .”
Evolutionary trajectory of SARS-CoV-2 and emerging variants
Singh, J., Pandit, P., McArthur, A. G., Banerjee, A., & Mossman, K. L. (2021) · Virology Journal · doi:10.1186/s12985-021-01633-w · Cited by 150
Fagre, A. C., Lewis, J., Eckley, M., Zhan, S., Rocha, S. M., Sexton, N. R., Burke, B. E., Geiss, B. J., Peersen, O. B., Bass, T. A., et al. (2021) · PLoS Pathogens · doi:10.1371/journal.ppat.1009585 · Cited by 145
SARS-CoV-2 and COVID-19: A genetic, epidemiological, and evolutionary perspective
Sironi, M., Hasnain, S. E., Rosenthal, B. M., Phan, T. G., Luciani, F., Shaw, M., Sallum, M. A. M., Mirhashemi, M. E., Morand, S., & Gónzález‐Candelas, F. (2020) · Infection Genetics and Evolution · doi:10.1016/j.meegid.2020.104384 · Cited by 145
Lost in deletion: The enigmatic ORF8 protein of SARS-CoV-2
Zinzula, L. (2020) · Biochemical and Biophysical Research Communications · doi:10.1016/j.bbrc.2020.10.045 · Cited by 136
“117 this well correlates with the proposed theory of SARS-CoV-2 originating from recombination events between horseshoe bat coronaviruses, as well as with the plausible role of pangolins as intermediate hosts that preceded SARS-CoV-2 spillover into human population [4,17].”
Binding and molecular basis of the bat coronavirus RaTG13 virus to ACE2 in humans and other species
Liu, K., Pan, X., Li, L., Yu, F., Zheng, A., Du, P., Han, P., Meng, Y., Zhang, Y., Wu, L., et al. (2021) · Cell · doi:10.1016/j.cell.2021.05.031 · Cited by 136
“3% amino acid identity in the receptor binding domain (RBD) (Boni et al., 2020; Wu et al., 2020a; Zhou et al., 2020b).”
InfluentialTang, T., Jaimes, J. A., Bidon, M. K., Straus, M. R., Daniel, S., & Whittaker, G. R. (2021) · ACS Infectious Diseases · doi:10.1021/acsinfecdis.0c00701 · Cited by 136
SARS-CoV-2 Infects Neurons and Induces Neuroinflammation in a Non-Human Primate Model of COVID-19
Beckman, D., Bonillas, A. C., Diniz, G. B., Ott, S. P., Roh, J. W., Elizaldi, S. R., Schmidt, B., Sammak, R. L., Rompay, K. K. A. V., Iyer, S. S., et al. (2022) · Cell Reports · doi:10.1016/j.celrep.2022.111573 · Cited by 132
Thompson, C. W., Phelps, K. L., Allard, M. W., Cook, J. A., Dunnum, J. L., Ferguson, A. W., Gelang, M., Khan, F. A. A., Paul, D. L., Reeder, D. M., et al. (2021) · mBio · doi:10.1128/mbio.02698-20 · Cited by 133
“SARS-CoV-2 (severe acute respiratory syndrome coronavirus 2) (3), the causative agent of COVID-19, has demonstrated that a previously unknown pathogen can emerge from wildlife species and threaten public health on a global scale within months.”
BackgroundInfluentialThe Role of Type I Interferons in the Pathogenesis and Treatment of COVID-19
Schreiber, G. (2020) · Frontiers in Immunology · doi:10.3389/fimmu.2020.595739 · Cited by 136
“Similar virus strains that circulate in bats in Hubei province in China may in the future cause further new zoonotic outbreaks (29).”
BackgroundAssessing the risks of SARS-CoV-2 in wildlife
Delahay, R. J., de la Fuente, J., Smith, G. C., Sharun, K., Snary, E., Giron, L. F., Nziza, J., Fooks, A. R., Brookes, S. M., Lean, F. Z. X., et al. (2021) · One Health Outlook · doi:10.1186/s42522-021-00039-6 · Cited by 128
Zoonotic disease and virome diversity in bats
Brussel, K. V., & Holmes, E. C. (2021) · Current Opinion in Virology · doi:10.1016/j.coviro.2021.12.008 · Cited by 128
Understanding the Role of SARS-CoV-2 ORF3a in Viral Pathogenesis and COVID-19
Zhang, J., Ejikemeuwa, A., Gerzanich, V., Nasr, M., Tang, Q., Simard, J. M., & Zhao, Y. (2022) · Frontiers in Microbiology · doi:10.3389/fmicb.2022.854567 · Cited by 138
Evolutionary dynamics of the SARS-CoV-2 ORF8 accessory gene
Pereira, F. (2020) · Infection Genetics and Evolution · doi:10.1016/j.meegid.2020.104525 · Cited by 123
“After the S gene, ORF8 stands out as being the most divergent gene when SARS-CoV-2 is compared with bat and pangolin coronaviruses (Boni et al., 2020).”
Recent advances in mRNA cancer vaccines: meeting challenges and embracing opportunities
Wang, B., Pei, J., Xu, S., Liu, J., & Yu, J. (2023) · Frontiers in Immunology · doi:10.3389/fimmu.2023.1246682 · Cited by 131
“Recently, the successful applications of COVID-19 global pandemic offered a great opportunity for messenger RNA (mRNA)vaccines in antitumor therapy (8 – 10).”
BackgroundLam, S. D., Bordin, N., Waman, V., Scholes, H. M., Ashford, P., Sen, N., van Dorp, L., Rauer, C., Dawson, N. L., Pang, C., et al. (2020) · Scientific Reports · doi:10.1038/s41598-020-71936-5 · Cited by 109
A clade of SARS-CoV-2 viruses associated with lower viral loads in patient upper airways
Lorenzo-Redondo, R., Nam, H. H., Roberts, S. C., Simons, L. M., Jennings, L. J., Qi, C., Achenbach, C. J., Hauser, A. R., Ison, M. G., Hultquist, J. F., et al. (2020) · EBioMedicine · doi:10.1016/j.ebiom.2020.103112 · Cited by 110
“6 £ 10 4 for the global tree, consistent with other studies [12,33].”
Reinfection in patients with COVID-19: a systematic review
Ren, X., Zhou, J., Guo, J., Hao, C., Zheng, M., Zhang, R., Huang, Q. Y., Yao, X., Li, R., & Jin, Y. (2022) · Global Health Research and Policy · doi:10.1186/s41256-022-00245-3 · Cited by 104
“Severe acute respiratory syndrome coronavirus 2 (SARSCoV-2) is a single-stranded positive-strand RNA virus that belongs to the Coronaviridae family [2, 3].”
BackgroundRisk of human‐to‐wildlife transmission of SARS‐CoV‐2
Gryseels, S., Bruyn, L. D., Gyselings, R., Calvignac‐Spencer, S., Leendertz, F. H., & Leirs, H. (2020) · Mammal Review · doi:10.1111/mam.12225 · Cited by 114
“taken place solely in its natural (horseshoe bat) reservoir or (partly) in another unidentified natural host [27].”
BackgroundInfluentialNeutralizing activity of Sputnik V vaccine sera against SARS-CoV-2 variants
Ikegame, S., Siddiquey, M. N. A., Hung, C.-T., Haas, G. D., Brambilla, L., Oguntuyo, K. Y., Kowdle, S., Chiu, H.-P., Stevens, C. S., Vilardo, A. E., et al. (2021) · Nature Communications · doi:10.1038/s41467-021-24909-9 · Cited by 141
Mapping genome variation of SARS-CoV-2 worldwide highlights the impact of COVID-19 super-spreadersPreprint
Gómez‐Carballa, A., Bello, X., Pardo‐Seco, J., Martinón‐Torres, F., & Salas, A. (2020) · Genome Research · doi:10.1101/gr.266221.120 · Cited by 109
“that could have existed in a time frame between 1948 and 1982 (Boni et al. 2020).”
BackgroundSun, D., Sang, Z., Kim, Y. J., Xiang, Y., Cohen, T., Belford, A. K., Huet, A., Conway, J. F., Sun, J., Taylor, D. J., et al. (2021) · Nature Communications · doi:10.1038/s41467-021-24963-3 · Cited by 96
Towards integrated surveillance-response systems for the prevention of future pandemics
Zinsstag, J., Utzinger, J., Probst-Hensch, N. M., Lv, S., & Zhou, X. (2020) · Infectious Diseases of Poverty · doi:10.1186/s40249-020-00757-5 · Cited by 108
“COVID-19 is suspected to result from animal-to-human transmission of SARS coronavirus 2 (SARS-CoV-2) with most close relation to a bat coronavirus [10, 11].”
BackgroundCOVID-19—lessons for zoonotic disease
Holmes, E. C. (2022) · Science · doi:10.1126/science.abn2222 · Cited by 102
Amoutzias, G. D., Nikolaidis, M., Tryfonopoulou, E., Chlichlia, K., Markoulatos, P., & Oliver, S. G. (2022) · Viruses · doi:10.3390/v14010078 · Cited by 99
“The feasibility of this scenario is evidenced by recent analyses of the role of recombination in Sarbecovirus evolution [52,76,111].”
BackgroundInfluentialNo evidence for distinct types in the evolution of SARS-CoV-2
MacLean, O. A., Orton, R. J., Singer, J. B., & Robertson, D. L. (2020) · Virus Evolution · doi:10.1093/ve/veaa034 · Cited by 96
Self-citationThe Emergence and Evolution of SARS-CoV-2
Holmes, E. C. (2024) · Annual Review of Virology · doi:10.1146/annurev-virology-093022-013037 · Cited by 96
Host Diversity and Origin of Zoonoses: The Ancient and the New
Recht, J., Schuenemann, V. J., & Sánchez-Villagra, M. R. (2020) · Animals · doi:10.3390/ani10091672 · Cited by 105
“A very recent phylogenetic dating study based on bioinformatic approaches strongly suggests that SARS-CoV-2 emerged directly, without an intermediate host, from the same horseshoe bat subgenus of SARS-like coronaviruses, with both SARS-CoV and SARS-CoV-2 diverging at the same time (40–70 years ago) from currently know…”
BackgroundGenomic sequencing effort for SARS-CoV-2 by country during the pandemic
Furuse, Y. (2020) · International Journal of Infectious Diseases · doi:10.1016/j.ijid.2020.12.034 · Cited by 90
“Collecting viral sequence data is also important for conducting an evolutionary analysis to infer the origin of the virus (Boni et al., 2020), detect mutations that may affect the pathogenicity and infectivity of the virus (Korber et al.”
BackgroundDynamically evolving novel overlapping gene as a factor in the SARS-CoV-2 pandemic
Nelson, C. W., Ardern, Z., Goldberg, T. L., Meng, C., Kuo, C.-H., Ludwig, C., Kolokotronis, S.-O., & Wei, X. (2020) · eLife · doi:10.7554/elife.59633 · Cited by 87
“Finally, given multiple recombination breakpoints in ORF3a (Boni et al., 2020) and the relative paucity of sequence data for viruses closely related to SARS-CoV-2, our analysis could not differentiate between convergence, recombination, or recurrent loss in the origin of ORF3d.”
BackgroundThe evolutionary history of ACE2 usage within the coronavirus subgenus Sarbecovirus
Wells, H., Letko, M. C., Lasso, G., Ssebide, B., Nziza, J., Byarugaba, D. K., Navarrete‐Macias, I., Liang, E., Cranfield, M. R., Han, B. A., et al. (2021) · Virus Evolution · doi:10.1093/ve/veab007 · Cited by 97
Cross-species recognition of SARS-CoV-2 to bat ACE2
Liu, K., Tan, S., Niu, S., Wang, J., Wu, L., Sun, H., Zhang, Y., Pan, X., Qu, X., Du, P., et al. (2020) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2020216118 · Cited by 89
“A growing body of research has identified that the binding capacity of ACE2 orthologs from different bat species to RBD of SARS-CoV or SARS-CoV-2 varied substantially, indicating the extensively diversified susceptibility of SARS-CoV and SARS-CoV-2 in different bat species (18, 25, 32, 33).”
Hosie, M. J., Epifano, I., Herder, V., Orton, R. J., Stevenson, A., Johnson, N., MacDonald, E., Dunbar, D., McDonald, M., Howie, F., et al. (2021) · Veterinary Record · doi:10.1002/vetr.247 · Cited by 91
Self-citationRandom-Forest-Bagging Broad Learning System With Applications for COVID-19 Pandemic
Zhan, C., Zheng, Y., Zhang, H., & Wen, Q. (2021) · IEEE Internet of Things Journal · doi:10.1109/jiot.2021.3066575 · Cited by 91
“The out-break of COVID-19 has progressed with a tremendous impact on the economic [3], social behavior [4], environment [5], climate [6], etc. Evolutionary virologist found bats may be the origins of SARS-CoV-2 [7], which has a long evolutionary history [8].”
BackgroundTian, J., Sun, J., Li, D., Wang, N., Wang, L., Zhang, C., Meng, X., Ji, X., Suchard, M. A., Zhang, X., et al. (2022) · Cell Reports · doi:10.1016/j.celrep.2022.110969 · Cited by 98
“Divergence dates between SARS-CoV-2 and other bat sarbecoviruses were estimated to be between 1948 and 1982, indicating that this lineage for SARSCoV-2 had been circulating in bats for several decades (Boni et al., 2020).”
Large‐scale genome sampling reveals unique immunity and metabolic adaptations in bats
Moreno-Santillán, D. D., Lama, T. M., Gutiérrez‐Guerrero, Y. T., Brown, A. M., Donat, P., Zhao, H., Rossiter, S. J., Yohe, L. R., Potter, J. H. T., Teeling, E. C., et al. (2021) · Molecular Ecology · doi:10.1111/mec.16027 · Cited by 87
“This probably includes the current SARS- CoV- 2 pandemic (Boni et al., 2020; Lau et al., 2020).”
BackgroundThe origins of COVID‐19 pandemic: A brief overview
Hao, Y., Wang, Y., Wang, M.-Y., Zhou, L., Shi, J., Cao, J., & Wang, D. (2022) · Transboundary and Emerging Diseases · doi:10.1111/tbed.14732 · Cited by 139
COVID-19: Time to exonerate the pangolin from the transmission of SARS-CoV-2 to humans
Frutos, R., Cobo, J. S., Chen, T., & Devaux, C. A. (2020) · Infection Genetics and Evolution · doi:10.1016/j.meegid.2020.104493 · Cited by 94
Metadata
- Authors
- Boni, M. F., Lemey, P., Jiang, X., Lam, T. T.-Y., Perry, B. W., Castoe, T. A., Rambaut, A., & Robertson, D. L.
- Affiliations
- Pennsylvania State University (US)
- KU Leuven (BE)
- Xi’an Jiaotong-Liverpool University (CN)
- University of Hong Kong (HK)
- The University of Texas at Arlington (US)
- University of Edinburgh (GB)
- MRC University of Glasgow Centre for Virus Research (GB)
- Corresponding author
- Boni, Maciej F.
- Lemey, Philippe
- Rambaut, Andrew
- Robertson, David L.
- ORCID
- Venue
- Nature Microbiology
- Published
- 2020-07-28
- Volume
- 5
- Issue
- 11
- Pages
- 1408-1417
- ISSN
- 2058-5276 (ISSN-L)
- OpenAlex
- W3046218112
- Cited by
- 1,014
- References
- 69
- Citations by year
- 2026: 23 · 2025: 46 · 2024: 79 · 2023: 118 · 2022: 240 · 2021: 327 · 2020: 178
- FWCI
- 28.54
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- Locations
- Nature Microbiology (publishedVersion)
- bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion, cc-by-nc)
- PubMed (publishedVersion)
- ENLIGHTEN (Jurnal Bimbingan dan Konseling Islam) (submittedVersion, cc-by-sa)
- Lirias (KU Leuven) (submittedVersion)
- Edinburgh Research Explorer (University of Edinburgh) (submittedVersion, other-oa)
- Funders
- Fonds Wetenschappelijk Onderzoek (G0D5117N, G0B9317N)
- Wellcome Trust (206298/Z/17/Z)
- RCUK | Medical Research Council (MC_UU_1201412)
- Topics
- Health Sciences › Medicine › Infectious Diseases › SARS-CoV-2 and COVID-19 Research
- Life Sciences › Agricultural and Biological Sciences › Animal Science and Zoology › Animal Virus Infections Studies
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Evolution and Genetic Dynamics
- Keywords
- SARS-CoV-2, sarbecoviruses, COVID-19 pandemic, viral recombination, bat reservoirs, pangolins, divergence time estimation, receptor-binding motif, ACE2 receptor, phylogenetic dating, viral emergence, genetic diversity, SARS-CoV
- MeSH terms
- Betacoronavirus, Angiotensin-Converting Enzyme 2, COVID-19, SARS-CoV-2, Animals, Bayes Theorem, China, Chiroptera, Humans, Peptidyl-Dipeptidase A, Phylogeny, Pneumonia, Viral, Recombination, Genetic, Genetic Variation, Genome, Viral, Coronavirus Infections, Evolution, Molecular, Pandemics
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