The DOI matches its Crossref record; 70 of the 70 references it cites were found in a registry or an index.
Evolutionary origins of the SARS-CoV-2 sarbecovirus lineage responsible for the COVID-19 pandemic
Open accessPublished version of a preprint
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.
Overview
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Integrity signals
Registry
Cites
Statistics
Verification
Claims the paper makes
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
In the citing paper
“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]
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
The self-check needs the paper’s full text.
Citations
Cites
RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies
Stamatakis, A. (2014) · Bioinformatics · doi:10.1093/bioinformatics/btu033 · Cited by 32,291 · cited 2×
“2.8 (Stamatakis, 2014, 2006), GTR+Γ model, 100 bootstrap replicates – was inferred for each BFR longer than 500nt.”
BackgroundMethodInfluentialIQ-TREE: A Fast and Effective Stochastic Algorithm for Estimating Maximum-Likelihood Phylogenies
Nguyen, L.-T., Schmidt, H. A., von Haeseler, A., & Minh, B. Q. (2015) · Molecular Biology and Evolution · doi:10.1093/molbev/msu300 · Cited by 26,046 · cited 1×
“The latter was reconstructed using IQTREE (Nguyen et al., 2014 ) under a General-Time Reversible (GTR) model with a discrete gamma distribution to model among-site rate variation.”
MethodA pneumonia outbreak associated with a new coronavirus of probable bat origin
Zhou, P., Yang, X.-L., Wang, X.-G., Hu, B., Zhang, L., Zhang, W., Si, H.-R., Zhu, Y., Li, B., Huang, C.-L., et al. (2020) · Nature · doi:10.1038/s41586-020-2012-7 · Cited by 21,969 · cited 11×
“…thought to be instrumental to SARS-CoV-2’s ability to infect humans – a polybasic cleavage site insertion in the Spike protein – has been seen in another close bat relative of the SARS-CoV-2 virus (Zhou et al., 2020a); however, the sequence is different and it is likely an independent event.”
BackgroundInfluentialEarly transmission dynamics in Wuhan, China, of novel coronavirus-infected pneumonia
Li, Q., Guan, X., Wu, P., Wang, X., Zhou, L., Tong, Y., Ren, R., Leung, K. S. M., Lau, E. H. Y., Wong, J. Y., et al. (2020) · New England Journal of Medicine · doi:10.1056/nejmoa2001316 · Cited by 16,575 · cited 8×
“In December 2019, a cluster of pneumonia cases epidemiologically linked to an open-air wet market in city of Wuhan (Hubei Province), China (Li et al., 2020a; Zhou et al., 2020b) led local health officials to issue an epidemiological alert to the Chinese Center for Disease Control and Prevention…”
BackgroundMethodInfluentialSelf-citationRAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models
Stamatakis, A. (2006) · Bioinformatics · doi:10.1093/bioinformatics/btl446 · Cited by 15,582 · cited 2×
“2.8 (Stamatakis, 2014, 2006), GTR+Γ model, 100 bootstrap replicates – was inferred for each BFR longer than 500nt.”
BackgroundMethodInfluentialLu, R., Zhao, X., Li, J., Niu, P., Yang, B., Wu, H., Wang, W., Song, H., Huang, B., Zhu, N., et al. (2020) · The Lancet · doi:10.1016/s0140-6736(20)30251-8 · Cited by 11,907 · cited 4×
“Across a large region of its genome, corresponding approximately to ORF1b , it did not cluster with any of the known bat coronaviruses indicating that recombination likely played a role in the evolutionary history of these viruses (Lu et al., 2020; Wu et al.”
BackgroundInfluentialA new coronavirus associated with human respiratory disease in China
Wu, F., Zhao, S., Yu, B., Chen, Y.-M., Wang, W., Song, Z.-G., Hu, Y., Tao, Z.-W., Tian, J.-H., Pei, Y.-Y., et al. (2020) · Nature · doi:10.1038/s41586-020-2008-3 · Cited by 11,688 · cited 4×
“When the first genome sequence of SARS-CoV-2, Wuhan-Hu-1, was released on January 10 2020 on Virological.org by a consortium led by Yong-Zhen Zhang (Wu et al., 2020a) it enabled the immediate analyses of its ancestry.”
BackgroundMethodInfluentialGorbalenya, A. E., Baker, S. C., Baric, R. S., de Groot, R. J., Drosten, C., Gulyaeva, A. A., Haagmans, B. L., Lauber, C., Leontovich, A. M., Neuman, B. W., et al. (2020) · Nature Microbiology · doi:10.1038/s41564-020-0695-z · Cited by 8,745 · cited 3×
“Next, we (1) collected all breakpoints into a single set, (2) complemented this set to generate a set of non-breakpoints, (3) grouped non-breakpoints into contiguous BFRs and (4) sorted these regions by length.”
BackgroundMethodThe proximal origin of SARS-CoV-2
Andersen, K. G., Rambaut, A., Lipkin, W. I., Holmes, E. C., & Garry, R. F. (2020) · Nature Medicine · doi:10.1038/s41591-020-0820-9 · Cited by 5,081 · cited 3×
“Influenza viruses reassort (Webster et al., 1992) but they do not undergo homologous recombination within RNA segments (Boni et al., 2008, 2010), meaning that ‘origins’ questions for influenza outbreaks can always be reduced to origins questions for each of influenza’s eight RNA segments.”
BackgroundMethodSelf-citationEvolution and ecology of influenza A viruses
Webster, R. G., Bean, W. J., Gorman, O. T., Chambers, T. M., & Kawaoka, Y. (1992) · Microbiological Reviews · doi:10.1128/mr.56.1.152-179.1992 · Cited by 4,739
Cited by
Sorted by citations
Items 701–800 of 997
Zaman, S., Sledzieski, S., Berger, B., Wu, Y.-C., & Bansal, M. S. (2022) · Journal of Computational Biology · doi:10.1089/cmb.2021.0507 · Cited by 5
Benazraf, A., & Arkin, I. T. (2022) · Protein Science · doi:10.1002/pro.4528 · Cited by 3
Som, A. (2022) · Bioinformation · doi:10.6026/97320630018951 · Cited by 3
Rooting and Dating Large SARS-CoV-2 Trees by Modeling Evolutionary Rate as a Function of Time
Xia, X. (2023) · Viruses · doi:10.3390/v15030684 · Cited by 3
Bhattacharyya, A., Anand, S., & Rai Shesh, N. (2023) · Annals of Public Health Reports · doi:10.36959/856/540 · Cited by 2
Eschatological events in Matthew 24:1–15 and COVID-19 pandemic in Nigeria
Ukeachusim, C. P. (2023) · Verbum et Ecclesia · doi:10.4102/ve.v44i1.2755 · Cited by 1
“Hence, some hold the position that COVID-19 is the result of genetic shuffling among known coronaviruses (Boni et al. 2020).”
BackgroundBats, Pathogen Diversity and Rabies in a Changing Neotropic Landscape
Rico‐Chávez, O., Flores-Pérez, N., Martínez-Pérez, K. U., Villalobos‐Segura, M. D. C., & Flores, R. Á. (2024) · Springer eBooks · doi:10.1007/978-3-031-50531-7_8 · Cited by 3
The human microbiota is a beneficial reservoir for SARS-CoV-2 mutations
Cao, B., Wang, X.-X., Yin, W., Gao, Z., & Xia, B.-Q. (2024) · mBio · doi:10.1128/mbio.03187-23 · Cited by 3
How to Survive COVID‐19 Even If the Vaccine Fails
Branch, A. D. (2020) · Hepatology Communications · doi:10.1002/hep4.1588 · Cited by 4
CORTICOSTEROIDS AND SECONDARY INFECTIONS: AN INSIGHT INTO CORONAVIRUS DISEASE-2019
Neela, B., Jakkula, S. V., MUJEEBUDDIN, & Gaur, R. P. (2021) · Asian Journal of Pharmaceutical and Clinical Research · doi:10.22159/ajpcr.2021.v14i1.40096 · Cited by 3
“In a recently published study, it has further established that sarbecovirus lineage is responsible for the pandemic of COVID-19 and concluded that the SARS-CoV-2 itself is not a recombinant of any sarbecoviruses detected to date and has been circulating unnoticed in bats for decades [10].”
BackgroundInvestigations into the Origin of SARS-CoV-2: An Update
Mitra, A. (2021) · Current Science · doi:10.18520/cs/v121/i1/77-84 · Cited by 3
Striking lineage diversity of severe acute respiratory syndrome coronavirus 2 from non-human sources
Muñoz, M., Patiño, L. H., Ballesteros, N., Castañeda, S., Luna, N., Noguera, L. A. D., Hernández‐Pereira, C. E., Shaban, M. V., Muñoz, S. A., Paniz‐Mondolfi, A., et al. (2021) · One Health · doi:10.1016/j.onehlt.2021.100363 · Cited by 3
“Such information would also contribute to the development of COVID-19 vaccines and the characterization of mutants that are likely to arise as the virus continues to travel from humans to animals to the environment, and vice versa [34].”
Simon, U. K. (2021) · Contributions from science education research · doi:10.1007/978-3-030-75297-2_14 · Cited by 3
The Pandemic of Tourism: How Tourism Has Become an Unsustainable Luxury
Hindley, C., Legrand, W., & Laeis, G. C. M. (2022) · Emerald Group Publishing Limited eBooks · doi:10.1108/978-1-83982-900-020211024 · Cited by 3
GenomeBits insight into omicron and delta variants of coronavirus pathogen
Canessa, E., & Tenze, L. (2022) · PLoS ONE · doi:10.1371/journal.pone.0271039 · Cited by 3
“These polymorphisms reveal the complex dynamic gene transfer-recombination events among lineages with potential to cause major human outbreaks [8].”
Background[Progress in source tracking of SARS-CoV-2].
He, W., & Chen, Q. (2020) · PubMed Central · doi:10.12122/j.issn.1673-4254.2020.12.22 · Cited by 3
Ansari, M. A., Alomary, M. N., Jamal, Q. M. S., Almoshari, Y., Salawi, A., Almahmoud, S. A., & Khan, J. (2022) · Current Pharmaceutical Design · doi:10.2174/1381612829666221019144259 · Cited by 3
Schrago, C. G., & Barzilai, L. P. (2021) · Genetics and Molecular Biology · doi:10.1590/1678-4685-gmb-2020-0254 · Cited by 3
“…for establishing transmission times is not applicable to the origin of the SARS-CoV-2 pandemic, because its closest known sister lineage, the RaTG13 sequence is evolutionarily distant, artificially increasing the lower bound for the age of the common ancestor (stem node) (Boni et al ., 2020).”
InfluentialEcology and Evolution of Betacoronaviruses
Rodríguez‐Román, E., & Gibbs, A. J. (2021) · Advances in experimental medicine and biology · doi:10.1007/978-3-030-63761-3_3 · Cited by 3
WITHDRAWNPreprint
Datta, S. (2021) · ChemRxiv · doi:10.26434/chemrxiv.13102877.v21 · Cited by 3
From partial to whole genome imputation of SARS-CoV-2 for epidemiological surveillancePreprint
Ortuño, F. M., Loucera, C., Casimiro‐Soriguer, C. S., Lepe, J. A., Camacho‐Martínez, P., Díaz, L. M., de Salazar, A., Chueca, N., García, F., Pérez-Florido, J., et al. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.04.13.439668 · Cited by 3
“Whole genome sequencing (WGS) has been successfully used for classification [2], studying transmission dynamics [3], and evaluating global and regional patterns of pandemic spread [4].”
MethodCOVID-19 Serology in New York State Using a Multiplex Microsphere ImmunoassayPreprint
Hunt, D., Yates, J. L., Kulas, K. E., Carson, K. J., Lamson, T., Demarest, V. L., Furuya, A. K. M., Howard, K. M., Marchewka, M. J., Stone, R., et al. (2021) · medRxiv · doi:10.1101/2021.05.12.21257125 · Cited by 2
Jolma, E. R., Gibson, L., Suu‐Ire, R., Fleischer, G., Asumah, S., Languon, S., Restif, O., Wood, J. L. N., & Cunningham, A. A. (2021) · Preprints.org · doi:10.20944/preprints202107.0467.v1 · Cited by 3
Zaman, S., Sledzieski, S., Berger, B., Wu, Y.-C., & Bansal, M. S. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.08.12.456131 · Cited by 3
Haney, J., Víjayakríshnan, S., Streetley, J., Dee, K., Goldfarb, D. M., Clarke, M., Mullin, M., Carter, S. D., Bhella, D., & Murcia, P. R. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.08.16.456460 · Cited by 3
“Direct interactions between viruses within coinfected cells can 41 result in changes to viral progenies, including -but not limited to- pseudotyping (incorporation 42 of surface proteins from a different virus) (8-10) or genomic rearrangements, which may 43 generate novel strains with pandemic potential such as SARS-C…”
BackgroundUse of eVLP-based vaccine candidates to broaden immunity against SARS-CoV-2 variantsPreprint
Bozic, J., Ahmed, T., Ontsouka, B., Fluckiger, A.-C., Diress, A., Berthoud, T. K., Yuan, X., Yang, L., Díaz‐Mitoma, F., Anderson, D. E., et al. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.09.28.462109 · Cited by 3
“Indeed, all approved vaccines have been designed against the ancestral SARS-COV-2 virus that is no longer circulating but has been replaced by variants containing mutations which are enabling escape from nAbs induced against the ancestral strain (Berio et al. 2020; Boni et al, 2020).”
BackgroundMicrobial Ecology and Evolution Are Essential for Understanding Pandemics
Frederickson, M. E., & Reese, A. T. (2021) · mBio · doi:10.1128/mbio.02144-21 · Cited by 3
“lineage “spilled over” into humans (16, 17) and to trace its spread around the globe (1,”
BackgroundLARGE-SCALE GENOME SAMPLING REVEALS UNIQUE IMMUNITY AND METABOLIC ADAPTATIONS IN BATSPreprint
Moreno-Santillán, D. D., Lama, T. M., Guerrero, Y. G., Brown, A. M., Donat, P., Zhao, H., Rossiter, S. J., Yohe, L. R., Potter, J. H. T., Teeling, E. C., et al. (2021) · doi:10.22541/au.160977727.76870866/v3 · Cited by 4
Sarbecoviruses of British Horseshoe Bats; Sequence Variation and EpidemiologyPreprint
Apaa, T., Withers, A. J., Staley, C. E., Blanchard, A. M., Bennett, M., Bremner‐Harrison, S., Chadwick, E. A., Hailer, F., Harrison, S. W. R., Loose, M., et al. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.02.14.528476 · Cited by 3
Genetic Recombination Sites Away from the Insertion/Deletion Hotspots in SARS-Related Coronaviruses
Akaishi, T., Fujiwara, K., & Ishii, T. (2022) · The Tohoku Journal of Experimental Medicine · doi:10.1620/tjem.2022.j093 · Cited by 2
Genomics and infectious diseases: Lessons learnt from the COVID-19 pandemic
Jolly, B., & Scaria, V. (2024) · Elsevier eBooks · doi:10.1016/b978-0-323-91799-5.00020-6 · Cited by 2
Ye, R., Zhao, J.-Q., Xie, H., Zhao, L., Gong, C., Wang, Z., Yue, N., Xia, L., Song, K., Dong, B., et al. (2025) · Emerging Microbes & Infections · doi:10.1080/22221751.2025.2494704 · Cited by 2
“The global COVID-19 pandemic further emphasizes the urgent need for identifying the etiology of acute respiratory tract infections for early public health interventions and effective clinical management [4].”
The Phylodynamic Threshold of Measurably Evolving Populations
Weber, A., Kende, J., González, C. D., Översti, S., & Duchêne, S. (2026) · Molecular Biology and Evolution · doi:10.1093/molbev/msag133 · Cited by 1
Babar, Q. A., Saeed, A., Bashir, M., Niaz, K., Rafiq, M., Waqas, M. Y., Tahir, M. F., Aqib, A. I., Dilawar, M., & Nisar, M. F. (2025) · BENTHAM SCIENCE PUBLISHERS eBooks · doi:10.2174/9789815322194125010006 · Cited by 2
Pärssinen, M., Sillanpää, I., & Kotila, M. (2021) · American Journal of Epidemiology and Infectious Disease · doi:10.12691/ajeid-9-1-4 · Cited by 2
COVID-19 as a zoonotic infection
Куличенко, А. Н., Maletskaya, O. V., Саркисян, Н. С., & Сергеевна, В. А. (2021) · Russian Journal of Infection and Immunity · doi:10.15789/2220-7619-caa-1621 · Cited by 2
Abdelsattar, A. S., El-Awadly, Z. M., Abdelgawad, M., Mahmoud, F., Allam, S. A., & Helal, M. A. (2021) · The Open COVID Journal · doi:10.2174/2666958702101010216 · Cited by 2
Morais, P., Trovão, N. S., Abecasis, A. B., & Parreira, R. (2022) · Virus Research · doi:10.1016/j.virusres.2022.198727 · Cited by 2
Allaka, T. R., Katari, N. K., & Jonnalagadda, S. B. (2022) · Physical Sciences Reviews · doi:10.1515/psr-2021-0089 · Cited by 2
INQUIETAÇÕES SOBRE A NATUREZA: ENTRE UTOPIAS E DISTOPIAS ANTRÓPICAS
Barbosa, C. S. (2022) · Caminhos de Geografia · doi:10.14393/rcg238658328 · Cited by 2
Reggiani, A., Rugna, G., & Bonilauri, P. (2022) · Frontiers in Veterinary Science · doi:10.3389/fvets.2022.1085613 · Cited by 2
“Despite its origin has not been completely understood, it has been demonstrated that this virus descends from bat coronaviruses and could have been transmitted to humans via a still unknown intermediate host (14, 15).”
BackgroundUnraveling COVID-19: A Global Health Crisis and Ongoing Research
Alam, O., Ahmad, M., Qureshi, M., Gul, M., Khan, N. A., Samad, A., Khan, Z., Atiq, S., Iqbal, A., & Ullah, A. (2023) · Futuristic Biotechnology · doi:10.54393/fbt.v3i02.47 · Cited by 2
Therapeutic potential of Lonicerae japonicae flos against emerging respiratory viral infections
Yang, J., Xia, B., Han, Y., Wang, Y., Bi, Y., & Zhang, Y. (2024) · Pharmacological Research - Modern Chinese Medicine · doi:10.1016/j.prmcm.2024.100362 · Cited by 2
Characterization of a pangolin SARS-CoV-2-related virus isolate that uses the human ACE2 receptor
Xia, L., Wang, X., Cui, X.-M., Zhang, Y.-M., Wang, Z., Li, E., Fan, C., Song, K., Li, Y.-G., Ye, R., et al. (2024) · Science China Life Sciences · doi:10.1007/s11427-023-2484-x · Cited by 3
“MT072864) (Lam et al., 2020).”
InfluentialApoorva, Shukla, A., Kumar, A., Singh, S. P., & Singh, S. K. (2025) · The Microbe · doi:10.1016/j.microb.2025.100237 · Cited by 2
COVID-19 vaccination induces cross-neutralisation of sarbecoviruses related to SARS-CoV-2
West, G. E., Morse, R. B., Sievers, B. L., Abdullahi, A., Kamelian, K., Cheng, M. T. K., Bradley, J. R., Baker, S., Graves, B., Stark, H., et al. (2026) · npj Vaccines · doi:10.1038/s41541-026-01469-x · Cited by 2
Theories of the origin of SARS-CoV-2 in the light of its continuing evolution
Débarre, F., & Hensel, Z. (2025) · Comptes Rendus Biologies · doi:10.5802/crbiol.183 · Cited by 2
Upham, N. S., Agosti, D., Poelen, J. H., Penev, L., Paul, D. L., Reeder, D. M., Simmons, N. B., Csorba, G., Groom, Q., Dimitrova, M., et al. (2020) · Biodiversity Information Science and Standards · doi:10.3897/biss.4.59199 · Cited by 2
Тихонов, Д., & Vladimirtsev, V. A. (2020) · Siberian Research · doi:10.33384/26587270.2020.04.02.06e · Cited by 2
Are new genome variants detected in SARS-CoV-2 expected considering population dynamics in viruses?
Ergören, M. Ç., Tulay, P., & Dündar, M. (2021) · The EuroBiotech Journal · doi:10.2478/ebtj-2021-0001 · Cited by 2
“Its specific binding ability to human Angiotensin-Converting Enzyme 2 (ACE2) receptor shows to be shared ancestral trait with two severe acute respiratory syndrome (SARS)-like bat coronaviruses, bat-SL-CoVZC45 and bat-CoVZXC2, but there is no direct evidence and molecular mechanisms still need to be resolved (3, 4).”
BackgroundLe virus SARS-CoV-2 n’a pas « d’origine »
Frutos, R., Gavotte, L., & Devaux, C. A. (2022) · médecine/sciences · doi:10.1051/medsci/2022079 · Cited by 2
COVID-19 in Missouri 2020-2021: A Perspective on Origins, Spread & Controversies, Part I.
McKinsey, D. S., McKinsey, J. P., Hampson, N. B., & Enriquez, M. (2022) · PubMed Central · Cited by 2
Jones, A., Massey, S. E., Zhang, D., Deigin, Y., & Quay, S. C. (2022) · Applied Microbiology · doi:10.3390/applmicrobiol2040068 · Cited by 2
“Neither (possible) recombination regions 1 or 2 were previously identified in other SARS2r-CoV genomes [14,59,60].”
A ciência como um processo: lições epistemológicas da pandemia
Guzzo, G. B., & Dall’Alba, G. (2021) · ACTIO Docência em Ciências · doi:10.3895/actio.v6n3.13133 · Cited by 2
The Potential of Natural Products in the Management of COVID-19
Saber, M. M., Salama, M. M., & Badary, O. A. (2024) · Advances in experimental medicine and biology · doi:10.1007/978-3-031-61939-7_12 · Cited by 2
How Trustworthy Are the Genomic Sequences of SARS-CoV-2 in GenBank?
Xia, X. (2024) · Microorganisms · doi:10.3390/microorganisms12112187 · Cited by 2
Genome data artifacts and functional studies of deletion repair in the BA.1 SARS-CoV-2 spike protein
Álvarez-Herrera, M., Ruiz-Rodríguez, P., Navarro-Domínguez, B. M., Zulaica, J., Grau, B., Bracho, M. A., Guerreiro, M., Aguilar-Gallardo, C., Gónzález‐Candelas, F., Comas, I., et al. (2025) · Virus Evolution · doi:10.1093/ve/veaf015 · Cited by 2
“Recombination might have played a role in the evolution of SARS-CoV-2 from the ancestral coronavirus before diversification in humans (Boni et al. 2020, Andersen et al. 2020, Kirtipal et al. 2020, MacLean et al. 2021, Wells et al. 2021), and studies using focused datasets have also pointed to…”
A theory for curing the diseases of modernity
Krakauer, J. W. (2020) · Current Biology · doi:10.1016/j.cub.2020.10.055 · Cited by 3
“Most recent evidence suggests that SARSCoV-2, the coronavirus that causes COVID-19, came from bats in the Yunnan province in China [2].”
Backgroundda Costa, A. P. A., da Rocha, V. L., Ramos, R. R., Neto, J. M. P., da Silva, W. R., Pereira, L. L. V., Ramos, K., Marques, T. C. N., Filho, I. J. Z., & Lozano, A. W. (2021) · Coronaviruses · doi:10.2174/2666796702666210118153843 · Cited by 2
A year after the start of COVID-19
Camacho-Ortíz, A. (2021) · Medicina Universitaria · doi:10.24875/rmu.m21000056 · Cited by 2
Fulci, V., Carissimi, C., & Laudadio, I. (2021) · OMICS A Journal of Integrative Biology · doi:10.1089/omi.2021.0058 · Cited by 3
“No-tably, both H1N1 and SARS-CoV-2 viral infections have a zoonotic origin (Andersen et al., 2020; Boni et al., 2020; Smith et al., 2009), have indeed been transmitted to humans from wild or domestic animals through direct contacts or indirectly through exposure to the urine or feces of infected…”
Motule, A. S. (2021) · International Journal for Research in Applied Science and Engineering Technology · doi:10.22214/ijraset.2021.37739 · Cited by 2
Los murciélagos y la COVID-19, una injusta historia
Galindo-González, J. R., & Medellín, R. A. (2021) · CIENCIA ergo sum · doi:10.30878/ces.v28n2a11 · Cited by 2
Detailed phylogenetic analysis of SARS-CoV-2 reveals latent capacity to bind human ACE2 receptorPreprint
Brintnell, E., Gupta, M., & Anderson, D. W. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.06.22.165787 · Cited by 2
Structural Variants in SARS-CoV-2 Occur at Template-Switching HotspotsPreprint
Chrisman, B., Paskov, K. M., Stockham, N., Tabatabaei, K., Jung, J.-Y., Washington, P. Y., Varma, M., Sun, M., Maleki, S., & Wall, D. P. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.09.01.278952 · Cited by 2
“Analysis on the sarbecovirus recombinant origins of SARS-CoV-2 identified possible recombination breakpoints at 1684, 3046, 9237, 11885, 21753, 22773 and 24628.(3).”
BackgroundFarah, I., Lalli, G., Baker, D. J., & Schumacher, A. (2020) · medRxiv · doi:10.1101/2020.09.28.20203257 · Cited by 2
Rahalkar, M. C., & Bahulikar, R. A. (2020) · Preprints.org · doi:10.20944/preprints202008.0205.v3 · Cited by 2
SARS-CoV-2 and Zoonotic Preparedness: Unknown Knowns?
Irving, A. T., & Welburn, S. C. (2021) · Infectious Microbes & Diseases · doi:10.1097/im9.0000000000000051 · Cited by 2
“While most CoVs exist in animals, SARS-CoV-2 is one of seven CoVs known to actively infect humans, all having likely derived from animal hosts.(2,3) The majority of CoV infections result in mild infection (eg, 229E) though MERS-CoV, SARS-CoV and SARSCoV-2 may result in severe morbidity and mortality.”
BackgroundHuman endemic coronavirus emergence in the context of past and recent zoonotic outbreaksPreprint
Forni, D., Cagliani, R., Arrigoni, F., Benvenuti, M., Pontremoli, C., Pozzoli, U., Clerici, M., GIOIA, L. D., & Sironi, M. (2021) · Research Square · doi:10.21203/rs.3.rs-134999/v1 · Cited by 2
Unlocking capacities of viral genomics for the COVID-19 pandemic responsePreprint
Knyazev, S., Chhugani, K., Sarwal, V., Ayyala, R., Singh, H., Karthikeyan, S., Deshpande, D., Comarova, Z., Lu, A., Porozov, Y. B., et al. (2021) · PubMed · doi:10.48550/arxiv.2104.14005 · Cited by 2
Self-citationEvolutionary trajectory of SARS-CoV-2 genomePreprint
Wang, A. (2021) · Research Square · doi:10.21203/rs.3.rs-1009010/v1 · Cited by 2
Considerations for an ethic of One Health : Towards a socially responsible zoonotic disease control
van Herten, J. (2021) · doi:10.18174/555123 · Cited by 4
Ijaz, M. K., Nims, R. W., Zhou, S. S., Whitehead, K., Sriniva, V., Kapes, T., Fanuel, S., Epstein, J. H., Daszak, P., Rubino, J. R., et al. (2020) · doi:10.21203/rs.3.rs-75158/v1 · Cited by 2
Yi, C., Sun, X., Lin, Y., Gu, C., Ding, L., Lu, X. P., Yang, Z. Y., Zhang, Y., Ma, L., Gu, W., et al. (2021) · Research Square · doi:10.21203/rs.3.rs-497595/v1 · Cited by 2
Chances of finding COVID-virus ancestor ‘almost nil’, say virologists
Mallapaty, S. (2022) · Nature · doi:10.1038/d41586-022-03611-w · Cited by 2
Rosenblatt, E., Cook, J. D., DiRenzo, G. V., Grant, E. H. C., Arce, F., Pepin, K. M., Rudolph, F. J., Runge, M. C., Shriner, S. A., Walsh, D. P., et al. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.08.30.555493 · Cited by 2
Veltri, P., Giancotti, R., Lomoio, U., Puccio, B., Tradigo, G., Guzzi, P. H., Vizza, P., & Torti, C. (2023) · doi:10.22541/au.169570833.31408402/v1 · Cited by 2
Hetland, M. A. K., Winkler, M. A., Kaspersen, H. P., Håkonsholm, F., Bakksjø, R., Bernhoff, E., Delgado‐Blas, J. F., Brisse, S., Correia, A., Fostervold, A., et al. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.09.11.612360 · Cited by 2
On the Controversies Surrounding the Lab-Leak Theory of COVID-19
Yoo, J. (2025) · Journal of Korean Medical Science · doi:10.3346/jkms.2025.40.e153 · Cited by 2
Asner, G. P. (2020) · The Conversation · doi:10.64628/aai.cfsp7grfj · Cited by 2
Ogunbode, T. O., Esan, V. I., Oyebamiji, V. O., Olatubi, I. V., & Ogunlaran, O. M. (2024) · Environmental Health Insights · doi:10.1177/11786302241266051 · Cited by 1
“12,13 This report has aggravated bitterness against bats despite findings identifying man as the main cause of his own public health challenges.”
BackgroundTimeline of the pandemic: epidemiology, global spread, variants and waves
Liu, Y.-X., Luo, J.-Y., Liu, R., Li, Y., & Guan, W. (2024) · European Respiratory Society eBooks · doi:10.1183/2312508x.10018823 · Cited by 1
Heidarzadeh, H., & Asad, S. H. (2024) · Plasmonics · doi:10.1007/s11468-024-02633-6 · Cited by 1
Virology Applications to the COVID-19 Pandemic
Kostaki, E. G. (2025) · Life · doi:10.3390/life15020247 · Cited by 1
Paiva, D. D. A., Yotoko, K. S. C., Zarzar, C. A., de Paiva, J. T., Siqueira, D. P., Rodrigues, G., de Souza, L. P. M., & Sáfadi, T. (2025) · Caderno Pedagógico · doi:10.54033/cadpedv22n8-177 · Cited by 1
In vivo antibody diversification targeting a conserved coronavirus epitope
Nair, U., Feng, Z., Akauliya, M., Esposito, A., Crain, C. R., Lamperti, E. D., Prum, T., Warner, J. E., Madungwe, L., Dale, G. A., et al. (2025) · The Journal of Experimental Medicine · doi:10.1084/jem.20241563 · Cited by 1
Chidoti, V., Loisier, A., Zinyakasa, T., Bourgarel, M., Pinarello, V., Matope, G., Chiweshe, N., Missé, D., Mwandiringana, E., Nys, H. M. D., et al. (2025) · BMC Genomics · doi:10.1186/s12864-025-12015-9 · Cited by 1
“Coronaviruses' envelope spike (S) protein facilitates virus entry into host cells by binding to the cellular receptor angiotensin-converting enzyme 2 (ACE2) in humans [41–43].”
Superior RdRp Function Drives the Dominance of Prevalent GI.3 Norovirus Lineages
Lu, Q., Du, H., Jiang, X., Zeng, B., Li, T., & Dai, Y.-C. (2025) · Microorganisms · doi:10.3390/microorganisms14010011 · Cited by 1
“3[P13] strains in Asia versus Europe/America, suggests that following recombination, viral subpopulations evolve independently under different local pressures, forming transmission networks with regional characteristics [34,35].”
Self-citationSARS-CoV-2 and Emerging Foodborne Pathogens: Intriguing Commonalities and Obvious Differences
Abdelhamid, A. G., Faraone, J. N., Evans, J. P., Liu, S., & Yousef, A. E. (2022) · Pathogens · doi:10.3390/pathogens11080837 · Cited by 1
“Four genomic regions in SARS-CoV-2 have undergone recombination, with three of these regions showing similarity to those in bat coronavirus RaTG13 [40].”
BackgroundResultInfluentialAdvances in Diagnosis and Treatment for SARS-CoV-2 Variants
Akhter, N., Sana, S., Ahsan, M. A., Siddique, Z., Huraira, A., & Sana, S. (2023) · Infectious diseases · doi:10.5772/intechopen.107846 · Cited by 1
Identifying animal reservoirs during an epidemic
Salkeld, D., Hopkins, S. R., & Hayman, D. T. S. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198825920.003.0008 · Cited by 2
Causes and Consequences of Coronavirus Spike Protein Variability
Zech, F., Jung, C., Jacob, T., & Kirchhoff, F. (2024) · Viruses · doi:10.3390/v16020177
Dixson, J. D., Vumma, L., & Azad, R. K. (2024) · Microorganisms · doi:10.3390/microorganisms12102021 · Cited by 1
Characterization and evolutionary history of novel SARS-CoV-2-related viruses in bats from Cambodia
Ou, T. P., Guillebaud, J., Baidaliuk, A., Tum, S., Cheang, D., Delaune, D., Prot, M., Bruder, E., Machado, R. R. G., Pum, L., et al. (2026) · Nature Communications · doi:10.1038/s41467-026-75954-1 · Cited by 1
“Ourestimatesarewithin the range of the mean estimates previously reported for sarbecov-iruses using partial (non-recombinant regions) genome data 25 , and comparable to estimates obtained for SARS-CoV-2 26 or MERS-CoV 18 .”
InfluentialReduction of extracellular vimentin in blood provides protection against SARS-CoV-2 infection
Kim, H.-M., Wang, M., Zhai, C., Kim, S., Park, J., & Hong, S. (2025) · Virulence · doi:10.1080/21505594.2025.2568052 · Cited by 5
“However, none of these zoonotic virus genomes exhibit more than 98% similarity to that of SARS-CoV-2 [6–8], suggesting that these SARS-CoV-2-related coronaviruses are not direct ancestor of SARS-CoV-2.”
Тихонов, Д., & Vladimirtsev, V. A. (2020) · Siberian Research · doi:10.33384/26587270.2020.04.02.01r · Cited by 1
Another Time of Renewal : Pandemics, Power, and Knowledge
Kijima, J. (2020) · Institutional Repositories DataBase (IRDB) · doi:10.15068/00162399 · Cited by 1
Are pandemics associated with intensive livestock production?
Alvseike, O. A., Tollersrud, T., & Blagojević, B. (2021) · Veterinarski glasnik · doi:10.2298/vetgl201217002a · Cited by 1
Long-Term Survivors of Breast Cancer: A Growing Population
Bilani, N., Elimimian, E. B., Elson, L. C., Liang, H., & Nahleh, Z. A. (2021) · IntechOpen eBooks · doi:10.5772/intechopen.95798 · Cited by 1
“Genomic sequencing and phylogenetic analysis indicated that the coronavirus that causes COVID-19 is of the same subgenus as the severe acute respiratory syndrome (SARS) virus [35, 36].”
BackgroundORIGIN OF SARS-COV-2:NATURAL O SELECTION OR ADAPTATIVE SELECTION?
Martínez, P. M. (2021) · REBIOL · doi:10.17268/rebiol.2020.40.02.14
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
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review