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Novel Bat Alphacoronaviruses in Southern China Support Chinese Horseshoe Bats as an Important Reservoir for Potential Novel Coronaviruses

Identifieur interne : 001617 ( Pmc/Curation ); précédent : 001616; suivant : 001618

Novel Bat Alphacoronaviruses in Southern China Support Chinese Horseshoe Bats as an Important Reservoir for Potential Novel Coronaviruses

Auteurs : Susanna K. P. Lau [République populaire de Chine] ; Antonio C. P. Wong ; Libao Zhang ; Hayes K. H. Luk ; Jamie S. L. Kwok ; Syed S. Ahmed ; Jian-Piao Cai ; Pyrear S. H. Zhao ; Jade L. L. Teng [République populaire de Chine] ; Stephen K. W. Tsui ; Kwok-Yung Yuen [République populaire de Chine] ; Patrick C. Y. Woo [République populaire de Chine]

Source :

RBID : PMC:6563315

Abstract

While bats are increasingly recognized as a source of coronavirus epidemics, the diversity and emergence potential of bat coronaviruses remains to be fully understood. Among 1779 bat samples collected in China, diverse coronaviruses were detected in 32 samples from five different bat species by RT-PCR. Two novel alphacoronaviruses, Rhinolophus sinicus bat coronavirus HKU32 (Rs-BatCoV HKU32) and Tylonycteris robustula bat coronavirus HKU33 (Tr-BatCoV HKU33), were discovered from Chinese horseshoe bats in Hong Kong and greater bamboo bats in Guizhou Province, respectively. Genome analyses showed that Rs-BatCoV HKU32 is closely related to BatCoV HKU10 and related viruses from diverse bat families, whereas Tr-BatCoV HKU33 is closely related to BtNv-AlphaCoV and similar viruses exclusively from bats of Vespertilionidae family. The close relatedness of Rs-BatCoV HKU32 to BatCoV HKU10 which was also detected in Pomona roundleaf bats from the same country park suggests that these viruses may have the tendency of infecting genetically distant bat populations of close geographical proximity with subsequent genetic divergence. Moreover, the presence of SARSr-CoV ORF7a-like protein in Rs-BatCoV HKU32 suggests a common evolutionary origin of this accessory protein with SARS-CoV, also from Chinese horseshoe bats, an apparent reservoir for coronavirus epidemics. The emergence potential of Rs-BatCoV HKU32 should be explored.


Url:
DOI: 10.3390/v11050423
PubMed: 31067830
PubMed Central: 6563315

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PMC:6563315

Le document en format XML

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<name sortKey="Tsui, Stephen K W" sort="Tsui, Stephen K W" uniqKey="Tsui S" first="Stephen K. W." last="Tsui">Stephen K. W. Tsui</name>
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<nlm:aff id="af6-viruses-11-00423">School of Biomedical Sciences, Faculty of Medicine, The Chinese University of Hong Kong, Shatin, Hong Kong, China;
<email>jamie_slk@link.cuhk.edu.hk</email>
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<name sortKey="Yuen, Kwok Yung" sort="Yuen, Kwok Yung" uniqKey="Yuen K" first="Kwok-Yung" last="Yuen">Kwok-Yung Yuen</name>
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<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
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<name sortKey="Woo, Patrick C Y" sort="Woo, Patrick C Y" uniqKey="Woo P" first="Patrick C. Y." last="Woo">Patrick C. Y. Woo</name>
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<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
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<affiliation wicri:level="1">
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<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Carol Yu Centre for Infection, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
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<nlm:aff id="af4-viruses-11-00423">Collaborative Innovation Centre for Diagnosis and Treatment of Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Collaborative Innovation Centre for Diagnosis and Treatment of Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
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<title xml:lang="en" level="a" type="main">Novel Bat Alphacoronaviruses in Southern China Support Chinese Horseshoe Bats as an Important Reservoir for Potential Novel Coronaviruses</title>
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<name sortKey="Lau, Susanna K P" sort="Lau, Susanna K P" uniqKey="Lau S" first="Susanna K. P." last="Lau">Susanna K. P. Lau</name>
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<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
<email>skplau@hku.hk</email>
(S.K.P.L.);
<email>antonwcp@connect.hku.hk</email>
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<email>caijuice@hku.hk</email>
(J.-P.C.);
<email>pyrear@126.com</email>
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</affiliation>
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<wicri:regionArea>State Key Laboratory of Emerging Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
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<affiliation wicri:level="1">
<nlm:aff id="af3-viruses-11-00423">Carol Yu Centre for Infection, The University of Hong Kong, Pokfulam, Hong Kong, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
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</affiliation>
</author>
<author>
<name sortKey="Wong, Antonio C P" sort="Wong, Antonio C P" uniqKey="Wong A" first="Antonio C. P." last="Wong">Antonio C. P. Wong</name>
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<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
<email>skplau@hku.hk</email>
(S.K.P.L.);
<email>antonwcp@connect.hku.hk</email>
(A.C.P.W);
<email>hkhluk@hku.hk</email>
(H.K.H.L.);
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(S.S.A.);
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(P.S.H.Z.);
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</affiliation>
</author>
<author>
<name sortKey="Zhang, Libao" sort="Zhang, Libao" uniqKey="Zhang L" first="Libao" last="Zhang">Libao Zhang</name>
<affiliation>
<nlm:aff id="af5-viruses-11-00423">Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Laboratory of Wild Animal Conservation and Utilization, Guangdong Institute of Applied Biological Resources, Guangzhou 510000, China;
<email>zhanglb@giabr.gd.cn</email>
</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Luk, Hayes K H" sort="Luk, Hayes K H" uniqKey="Luk H" first="Hayes K. H." last="Luk">Hayes K. H. Luk</name>
<affiliation>
<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
<email>skplau@hku.hk</email>
(S.K.P.L.);
<email>antonwcp@connect.hku.hk</email>
(A.C.P.W);
<email>hkhluk@hku.hk</email>
(H.K.H.L.);
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<author>
<name sortKey="Kwok, Jamie S L" sort="Kwok, Jamie S L" uniqKey="Kwok J" first="Jamie S. L." last="Kwok">Jamie S. L. Kwok</name>
<affiliation>
<nlm:aff id="af6-viruses-11-00423">School of Biomedical Sciences, Faculty of Medicine, The Chinese University of Hong Kong, Shatin, Hong Kong, China;
<email>jamie_slk@link.cuhk.edu.hk</email>
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<email>kwtsui@cuhk.edu.hk</email>
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</affiliation>
</author>
<author>
<name sortKey="Ahmed, Syed S" sort="Ahmed, Syed S" uniqKey="Ahmed S" first="Syed S." last="Ahmed">Syed S. Ahmed</name>
<affiliation>
<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
<email>skplau@hku.hk</email>
(S.K.P.L.);
<email>antonwcp@connect.hku.hk</email>
(A.C.P.W);
<email>hkhluk@hku.hk</email>
(H.K.H.L.);
<email>shakeel87@gmail.com</email>
(S.S.A.);
<email>caijuice@hku.hk</email>
(J.-P.C.);
<email>pyrear@126.com</email>
(P.S.H.Z.);
<email>llteng@hku.hk</email>
(J.L.L.T.)</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Cai, Jian Piao" sort="Cai, Jian Piao" uniqKey="Cai J" first="Jian-Piao" last="Cai">Jian-Piao Cai</name>
<affiliation>
<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
<email>skplau@hku.hk</email>
(S.K.P.L.);
<email>antonwcp@connect.hku.hk</email>
(A.C.P.W);
<email>hkhluk@hku.hk</email>
(H.K.H.L.);
<email>shakeel87@gmail.com</email>
(S.S.A.);
<email>caijuice@hku.hk</email>
(J.-P.C.);
<email>pyrear@126.com</email>
(P.S.H.Z.);
<email>llteng@hku.hk</email>
(J.L.L.T.)</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Zhao, Pyrear S H" sort="Zhao, Pyrear S H" uniqKey="Zhao P" first="Pyrear S. H." last="Zhao">Pyrear S. H. Zhao</name>
<affiliation>
<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
<email>skplau@hku.hk</email>
(S.K.P.L.);
<email>antonwcp@connect.hku.hk</email>
(A.C.P.W);
<email>hkhluk@hku.hk</email>
(H.K.H.L.);
<email>shakeel87@gmail.com</email>
(S.S.A.);
<email>caijuice@hku.hk</email>
(J.-P.C.);
<email>pyrear@126.com</email>
(P.S.H.Z.);
<email>llteng@hku.hk</email>
(J.L.L.T.)</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Teng, Jade L L" sort="Teng, Jade L L" uniqKey="Teng J" first="Jade L. L." last="Teng">Jade L. L. Teng</name>
<affiliation>
<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
<email>skplau@hku.hk</email>
(S.K.P.L.);
<email>antonwcp@connect.hku.hk</email>
(A.C.P.W);
<email>hkhluk@hku.hk</email>
(H.K.H.L.);
<email>shakeel87@gmail.com</email>
(S.S.A.);
<email>caijuice@hku.hk</email>
(J.-P.C.);
<email>pyrear@126.com</email>
(P.S.H.Z.);
<email>llteng@hku.hk</email>
(J.L.L.T.)</nlm:aff>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="af2-viruses-11-00423">State Key Laboratory of Emerging Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>State Key Laboratory of Emerging Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="af3-viruses-11-00423">Carol Yu Centre for Infection, The University of Hong Kong, Pokfulam, Hong Kong, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Carol Yu Centre for Infection, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
</affiliation>
</author>
<author>
<name sortKey="Tsui, Stephen K W" sort="Tsui, Stephen K W" uniqKey="Tsui S" first="Stephen K. W." last="Tsui">Stephen K. W. Tsui</name>
<affiliation>
<nlm:aff id="af6-viruses-11-00423">School of Biomedical Sciences, Faculty of Medicine, The Chinese University of Hong Kong, Shatin, Hong Kong, China;
<email>jamie_slk@link.cuhk.edu.hk</email>
(J.S.L.K.);
<email>kwtsui@cuhk.edu.hk</email>
(S.K.W.T.)</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Yuen, Kwok Yung" sort="Yuen, Kwok Yung" uniqKey="Yuen K" first="Kwok-Yung" last="Yuen">Kwok-Yung Yuen</name>
<affiliation>
<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
<email>skplau@hku.hk</email>
(S.K.P.L.);
<email>antonwcp@connect.hku.hk</email>
(A.C.P.W);
<email>hkhluk@hku.hk</email>
(H.K.H.L.);
<email>shakeel87@gmail.com</email>
(S.S.A.);
<email>caijuice@hku.hk</email>
(J.-P.C.);
<email>pyrear@126.com</email>
(P.S.H.Z.);
<email>llteng@hku.hk</email>
(J.L.L.T.)</nlm:aff>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="af2-viruses-11-00423">State Key Laboratory of Emerging Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>State Key Laboratory of Emerging Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="af3-viruses-11-00423">Carol Yu Centre for Infection, The University of Hong Kong, Pokfulam, Hong Kong, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Carol Yu Centre for Infection, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="af4-viruses-11-00423">Collaborative Innovation Centre for Diagnosis and Treatment of Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Collaborative Innovation Centre for Diagnosis and Treatment of Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
</affiliation>
</author>
<author>
<name sortKey="Woo, Patrick C Y" sort="Woo, Patrick C Y" uniqKey="Woo P" first="Patrick C. Y." last="Woo">Patrick C. Y. Woo</name>
<affiliation>
<nlm:aff id="af1-viruses-11-00423">Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
<email>skplau@hku.hk</email>
(S.K.P.L.);
<email>antonwcp@connect.hku.hk</email>
(A.C.P.W);
<email>hkhluk@hku.hk</email>
(H.K.H.L.);
<email>shakeel87@gmail.com</email>
(S.S.A.);
<email>caijuice@hku.hk</email>
(J.-P.C.);
<email>pyrear@126.com</email>
(P.S.H.Z.);
<email>llteng@hku.hk</email>
(J.L.L.T.)</nlm:aff>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="af2-viruses-11-00423">State Key Laboratory of Emerging Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>State Key Laboratory of Emerging Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="af3-viruses-11-00423">Carol Yu Centre for Infection, The University of Hong Kong, Pokfulam, Hong Kong, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Carol Yu Centre for Infection, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="af4-viruses-11-00423">Collaborative Innovation Centre for Diagnosis and Treatment of Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Collaborative Innovation Centre for Diagnosis and Treatment of Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong</wicri:regionArea>
</affiliation>
</author>
</analytic>
<series>
<title level="j">Viruses</title>
<idno type="eISSN">1999-4915</idno>
<imprint>
<date when="2019">2019</date>
</imprint>
</series>
</biblStruct>
</sourceDesc>
</fileDesc>
<profileDesc>
<textClass></textClass>
</profileDesc>
</teiHeader>
<front>
<div type="abstract" xml:lang="en">
<p>While bats are increasingly recognized as a source of coronavirus epidemics, the diversity and emergence potential of bat coronaviruses remains to be fully understood. Among 1779 bat samples collected in China, diverse coronaviruses were detected in 32 samples from five different bat species by RT-PCR. Two novel alphacoronaviruses,
<italic>Rhinolophus sinicus</italic>
bat coronavirus HKU32 (Rs-BatCoV HKU32) and
<italic>Tylonycteris robustula</italic>
bat coronavirus HKU33 (Tr-BatCoV HKU33), were discovered from Chinese horseshoe bats in Hong Kong and greater bamboo bats in Guizhou Province, respectively. Genome analyses showed that Rs-BatCoV HKU32 is closely related to BatCoV HKU10 and related viruses from diverse bat families, whereas Tr-BatCoV HKU33 is closely related to BtNv-AlphaCoV and similar viruses exclusively from bats of
<italic>Vespertilionidae</italic>
family. The close relatedness of Rs-BatCoV HKU32 to BatCoV HKU10 which was also detected in Pomona roundleaf bats from the same country park suggests that these viruses may have the tendency of infecting genetically distant bat populations of close geographical proximity with subsequent genetic divergence. Moreover, the presence of SARSr-CoV ORF7a-like protein in Rs-BatCoV HKU32 suggests a common evolutionary origin of this accessory protein with SARS-CoV, also from Chinese horseshoe bats, an apparent reservoir for coronavirus epidemics. The emergence potential of Rs-BatCoV HKU32 should be explored.</p>
</div>
</front>
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</TEI>
<pmc article-type="research-article">
<pmc-dir>properties open_access</pmc-dir>
<front>
<journal-meta>
<journal-id journal-id-type="nlm-ta">Viruses</journal-id>
<journal-id journal-id-type="iso-abbrev">Viruses</journal-id>
<journal-id journal-id-type="publisher-id">viruses</journal-id>
<journal-title-group>
<journal-title>Viruses</journal-title>
</journal-title-group>
<issn pub-type="epub">1999-4915</issn>
<publisher>
<publisher-name>MDPI</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="pmid">31067830</article-id>
<article-id pub-id-type="pmc">6563315</article-id>
<article-id pub-id-type="doi">10.3390/v11050423</article-id>
<article-id pub-id-type="publisher-id">viruses-11-00423</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Article</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Novel Bat Alphacoronaviruses in Southern China Support Chinese Horseshoe Bats as an Important Reservoir for Potential Novel Coronaviruses</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Lau</surname>
<given-names>Susanna K.P.</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00423">1</xref>
<xref ref-type="aff" rid="af2-viruses-11-00423">2</xref>
<xref ref-type="aff" rid="af3-viruses-11-00423">3</xref>
<xref ref-type="aff" rid="af4-viruses-11-00423">4</xref>
<xref ref-type="author-notes" rid="fn1-viruses-11-00423"></xref>
</contrib>
<contrib contrib-type="author">
<contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0002-5565-1121</contrib-id>
<name>
<surname>Wong</surname>
<given-names>Antonio C.P.</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00423">1</xref>
<xref ref-type="author-notes" rid="fn1-viruses-11-00423"></xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zhang</surname>
<given-names>Libao</given-names>
</name>
<xref ref-type="aff" rid="af5-viruses-11-00423">5</xref>
<xref ref-type="author-notes" rid="fn1-viruses-11-00423"></xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Luk</surname>
<given-names>Hayes K.H.</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00423">1</xref>
</contrib>
<contrib contrib-type="author">
<contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0002-7550-9851</contrib-id>
<name>
<surname>Kwok</surname>
<given-names>Jamie S. L.</given-names>
</name>
<xref ref-type="aff" rid="af6-viruses-11-00423">6</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ahmed</surname>
<given-names>Syed S.</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00423">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Cai</surname>
<given-names>Jian-Piao</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00423">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zhao</surname>
<given-names>Pyrear S.H.</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00423">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Teng</surname>
<given-names>Jade L.L.</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00423">1</xref>
<xref ref-type="aff" rid="af2-viruses-11-00423">2</xref>
<xref ref-type="aff" rid="af3-viruses-11-00423">3</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Tsui</surname>
<given-names>Stephen K.W.</given-names>
</name>
<xref ref-type="aff" rid="af6-viruses-11-00423">6</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Yuen</surname>
<given-names>Kwok-Yung</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00423">1</xref>
<xref ref-type="aff" rid="af2-viruses-11-00423">2</xref>
<xref ref-type="aff" rid="af3-viruses-11-00423">3</xref>
<xref ref-type="aff" rid="af4-viruses-11-00423">4</xref>
<xref rid="c1-viruses-11-00423" ref-type="corresp">*</xref>
</contrib>
<contrib contrib-type="author">
<contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0001-9401-1832</contrib-id>
<name>
<surname>Woo</surname>
<given-names>Patrick C. Y.</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00423">1</xref>
<xref ref-type="aff" rid="af2-viruses-11-00423">2</xref>
<xref ref-type="aff" rid="af3-viruses-11-00423">3</xref>
<xref ref-type="aff" rid="af4-viruses-11-00423">4</xref>
<xref rid="c1-viruses-11-00423" ref-type="corresp">*</xref>
</contrib>
</contrib-group>
<aff id="af1-viruses-11-00423">
<label>1</label>
Department of Microbiology, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Pokfulam, Hong Kong, China;
<email>skplau@hku.hk</email>
(S.K.P.L.);
<email>antonwcp@connect.hku.hk</email>
(A.C.P.W);
<email>hkhluk@hku.hk</email>
(H.K.H.L.);
<email>shakeel87@gmail.com</email>
(S.S.A.);
<email>caijuice@hku.hk</email>
(J.-P.C.);
<email>pyrear@126.com</email>
(P.S.H.Z.);
<email>llteng@hku.hk</email>
(J.L.L.T.)</aff>
<aff id="af2-viruses-11-00423">
<label>2</label>
State Key Laboratory of Emerging Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong, China</aff>
<aff id="af3-viruses-11-00423">
<label>3</label>
Carol Yu Centre for Infection, The University of Hong Kong, Pokfulam, Hong Kong, China</aff>
<aff id="af4-viruses-11-00423">
<label>4</label>
Collaborative Innovation Centre for Diagnosis and Treatment of Infectious Diseases, The University of Hong Kong, Pokfulam, Hong Kong, China</aff>
<aff id="af5-viruses-11-00423">
<label>5</label>
Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Laboratory of Wild Animal Conservation and Utilization, Guangdong Institute of Applied Biological Resources, Guangzhou 510000, China;
<email>zhanglb@giabr.gd.cn</email>
</aff>
<aff id="af6-viruses-11-00423">
<label>6</label>
School of Biomedical Sciences, Faculty of Medicine, The Chinese University of Hong Kong, Shatin, Hong Kong, China;
<email>jamie_slk@link.cuhk.edu.hk</email>
(J.S.L.K.);
<email>kwtsui@cuhk.edu.hk</email>
(S.K.W.T.)</aff>
<author-notes>
<corresp id="c1-viruses-11-00423">
<label>*</label>
Correspondence:
<email>kyyuen@hku.hk</email>
(K.-Y.Y.);
<email>pcywoo@hku.hk</email>
(P.C.Y.W.); Tel.: +852-2255-4892 (K.-Y.Y. & P.C.Y.W.); Fax: +852-2855-1241 (K.-Y.Y. & P.C.Y.W.)</corresp>
<fn id="fn1-viruses-11-00423">
<label></label>
<p>These authors contributed equally to this manuscript.</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>07</day>
<month>5</month>
<year>2019</year>
</pub-date>
<pub-date pub-type="collection">
<month>5</month>
<year>2019</year>
</pub-date>
<volume>11</volume>
<issue>5</issue>
<elocation-id>423</elocation-id>
<history>
<date date-type="received">
<day>15</day>
<month>4</month>
<year>2019</year>
</date>
<date date-type="accepted">
<day>06</day>
<month>5</month>
<year>2019</year>
</date>
</history>
<permissions>
<copyright-statement>© 2019 by the authors.</copyright-statement>
<copyright-year>2019</copyright-year>
<license license-type="open-access">
<license-p>Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (
<ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">http://creativecommons.org/licenses/by/4.0/</ext-link>
).</license-p>
</license>
</permissions>
<abstract>
<p>While bats are increasingly recognized as a source of coronavirus epidemics, the diversity and emergence potential of bat coronaviruses remains to be fully understood. Among 1779 bat samples collected in China, diverse coronaviruses were detected in 32 samples from five different bat species by RT-PCR. Two novel alphacoronaviruses,
<italic>Rhinolophus sinicus</italic>
bat coronavirus HKU32 (Rs-BatCoV HKU32) and
<italic>Tylonycteris robustula</italic>
bat coronavirus HKU33 (Tr-BatCoV HKU33), were discovered from Chinese horseshoe bats in Hong Kong and greater bamboo bats in Guizhou Province, respectively. Genome analyses showed that Rs-BatCoV HKU32 is closely related to BatCoV HKU10 and related viruses from diverse bat families, whereas Tr-BatCoV HKU33 is closely related to BtNv-AlphaCoV and similar viruses exclusively from bats of
<italic>Vespertilionidae</italic>
family. The close relatedness of Rs-BatCoV HKU32 to BatCoV HKU10 which was also detected in Pomona roundleaf bats from the same country park suggests that these viruses may have the tendency of infecting genetically distant bat populations of close geographical proximity with subsequent genetic divergence. Moreover, the presence of SARSr-CoV ORF7a-like protein in Rs-BatCoV HKU32 suggests a common evolutionary origin of this accessory protein with SARS-CoV, also from Chinese horseshoe bats, an apparent reservoir for coronavirus epidemics. The emergence potential of Rs-BatCoV HKU32 should be explored.</p>
</abstract>
<kwd-group>
<kwd>
<italic>Alphacoronavirus</italic>
</kwd>
<kwd>novel</kwd>
<kwd>bat</kwd>
<kwd>coronavirus</kwd>
<kwd>species</kwd>
<kwd>molecular</kwd>
<kwd>discovery</kwd>
<kwd>epidemics</kwd>
</kwd-group>
</article-meta>
</front>
<floats-group>
<fig id="viruses-11-00423-f001" orientation="portrait" position="float">
<label>Figure 1</label>
<caption>
<p>Map of southern China showing locations where bat coronaviruses were found. Black bat represents the location with bats positive for Hi-BatCoV HKU10; orange bat represents the location with bats positive for coronavirus (CoV) PREDICT CoV-37; blue bat represents the location with bats positive for Rs-BatCoV HKU32; green bat represents the location with bats positive for Tr-BatCoV HKU33; grey bat represents the location with bats positive for severe acute respiratory syndrome related (SARSr) BatCoV; yellow bat represents the location with bats positive for Ty-BatCoV HKU4. Provinces where samples were collected are in red font.</p>
</caption>
<graphic xlink:href="viruses-11-00423-g001"></graphic>
</fig>
<fig id="viruses-11-00423-f002" orientation="portrait" position="float">
<label>Figure 2</label>
<caption>
<p>Genome organizations of Rs-BatCoV HKU32 strain TLC28A and Tr-BatCoV HKU33 strain GZ151867. Genes for ORF 1a and 1b of Rs-BatCoV HKU32 strain TLC28A and Tr-BatCoV HKU33 strain GZ151867 are represented by red and orange boxes, respectively. Genes for spike protein (S), envelope protein (E), membrane protein (M) and nucleocapsid protein (N) are represented by blue boxes. Genes for putative accessory proteins are represented by yellow boxes.</p>
</caption>
<graphic xlink:href="viruses-11-00423-g002"></graphic>
</fig>
<fig id="viruses-11-00423-f003" orientation="portrait" position="float">
<label>Figure 3</label>
<caption>
<p>Phylogenetic analysis of ORF1ab amino acid sequences of Rs-BatCoV HKU32 strains TLC26A and 28A, Tr-BatCoV HKU33 strain GZ151867 and other alphaCoVs. ORF1ab tree was constructed by maximum likelihood method using LG + G + I + F substitution model. The bootstrap values are calculated from 1000 trees. Tree was rooted using corresponding sequence of Middle East respiratory syndrome (MERS)-CoV (GenBank accession number YP_009047202.1). All bootstrap values are shown. The scale bar represents 5 substitutions per site. Both Rs-BatCoV HKU32 and Tr-BatCoV HKU33 are labeled with red (triangle) and blue (circle), respectively. Corresponding viral bat hosts’ families are highlighted in different colors: Red,
<italic>Vespertilionidae</italic>
; blue,
<italic>Miniopteridae</italic>
; yellow,
<italic>Rhinolophidae</italic>
; purple,
<italic>Hipposideridae</italic>
; green,
<italic>Pteropodidae</italic>
.</p>
</caption>
<graphic xlink:href="viruses-11-00423-g003"></graphic>
</fig>
<fig id="viruses-11-00423-f004" orientation="portrait" position="float">
<label>Figure 4</label>
<caption>
<p>Phylogenetic analysis of S amino acid sequences of Rs-BatCoV HKU32 strains TLC26A and 28A, Tr-BatCoV HKU33 strain GZ151867 and other alphaCoVs. S tree was constructed by maximum likelihood method using WAG + G + I + F substitution model. The bootstrap values are calculated from 1000 trees. Tree was rooted using corresponding sequence of MERS-CoV (GenBank accession number YP_009047204.1). All bootstrap values are shown. The scale bar represents 2 substitutions per site. Both Rs-BatCoV HKU32 and Tr-BatCoV HKU33 are highlighted in red (triangle) and blue (circle), respectively. Corresponding viral hosts are shown on the right.</p>
</caption>
<graphic xlink:href="viruses-11-00423-g004"></graphic>
</fig>
<fig id="viruses-11-00423-f005" orientation="portrait" position="float">
<label>Figure 5</label>
<caption>
<p>Rs-BatCoV HKU32 strain TLC28A mRNA leader-body junction and flanking sequences. The subgenomic ORF10 mRNA sequence are shown in alignment with the leader and genomic sequences. Identical nucleotides between the leader sequence and subgenomic mRNA sequence are labeled in green. Identical nucleotides between genome and subgenomic mRNA sequence are labeled in blue. The putative TRS is labeled in bold type, in purple and is underlined. Start codon AUG is labeled in red.</p>
</caption>
<graphic xlink:href="viruses-11-00423-g005"></graphic>
</fig>
<table-wrap id="viruses-11-00423-t001" orientation="portrait" position="float">
<object-id pub-id-type="pii">viruses-11-00423-t001_Table 1</object-id>
<label>Table 1</label>
<caption>
<p>Detection of CoVs in different bat species by reverse transcription (RT)-polymerase chain reaction (PCR) of the 440-bp fragment of RNA-dependent RNA polymerase (RdRp) gene.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Scientific Name</th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Common Name</th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">No. of Bats Captured</th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">No. of Bats Positive for CoV / (%)</th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">CoV Detected</th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Sampling Location of Bats</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Cynopterus sphinx</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Greater short-nosed fruit bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">SWH</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Hipposideros armiger</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Great roundleaf bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">GZ</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Hipposideros larvatus</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Intermediate roundleaf bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">21</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">GZ</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Hipposideros pomona</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Pomona leaf-nosed bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">182</td>
<td align="left" valign="middle" rowspan="1" colspan="1">2 / (1.1)</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Hi-BatCoV HKU10</td>
<td align="left" valign="middle" rowspan="1" colspan="1">TLC13, GD</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Hypsugo pulveratus</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Chinese pipistrelle</td>
<td align="left" valign="middle" rowspan="1" colspan="1">2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">LMHP</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Miniopterus magnater</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Western bent-winged bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">SK01</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Miniopterus pusillus</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Small bent-wing bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">56</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">LMH, SWH, SK01</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Miniopterus schreibersii</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Common bent-wing bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">23</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">SK01</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Miniopterus filiginosus</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Eastern bent-wing bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">LMHP</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Myotis chinensis</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Large myotis</td>
<td align="left" valign="middle" rowspan="1" colspan="1">10</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">SK01, GZ</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Myotis ricketti</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Rickett’s big-footed bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">93</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1 / (1.1)</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Coronavirus PREDICT CoV-37</td>
<td align="left" valign="middle" rowspan="1" colspan="1">LMH01, SK01</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Nyctalus noctula</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Common noctule</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">YSO</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Pipistrellus abramus</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Japanese pipistrelle</td>
<td align="left" valign="middle" rowspan="1" colspan="1">6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">MPO, YSO, KKSH</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Pipistrellus tenuis</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Least pipistrelle</td>
<td align="left" valign="middle" rowspan="1" colspan="1">4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">KKSH, YSO, SWH, LMHP</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Rhinolophus affinis</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Intermediate horseshoe bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">76</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">TLC01, TLC13, SK01</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Rhinolophus pearsonii</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Pearson’s horseshoe bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">GDP</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Rhinolophus pusillus</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Least horseshoe bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">17</td>
<td align="left" valign="middle" rowspan="1" colspan="1">0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">-</td>
<td align="left" valign="middle" rowspan="1" colspan="1">TLC13</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Rhinolophus sinicus</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Chinese horseshoe bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">272</td>
<td align="left" valign="middle" rowspan="1" colspan="1">10 / (3.7)</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Rs-BatCoV HKU32 (7)
<break></break>
SARSr BatCoV (3)</td>
<td align="left" valign="middle" rowspan="1" colspan="1">TLC01, GDP</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">
<italic>Tylonycteris pachypus</italic>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Lesser bamboo bat</td>
<td align="left" valign="middle" rowspan="1" colspan="1">240</td>
<td align="left" valign="middle" rowspan="1" colspan="1">18 / (7.5)</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Ty-BatCoV HKU4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">WKT, PFL, SWH, TLC01, GZP</td>
</tr>
<tr>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">
<italic>Tylonycteris robustula</italic>
</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Greater bamboo bat</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">104</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 / (0.96)</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tr-BatCoV HKU33</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GZP</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>GD, Guangdong Province; GDP, Guangdong Province—Conghua City; GZ, Guizhou Province; GZP, Guizhou Province—Luodian County; KKSH, Kai Kuk Shue Ha, Luk Keng; LMH, Lin Ma Hang Lead Mine; LMHP, Lin Ma Hang Pool; MPO, Mai Po Nature Reserve; PFL, Pok Fu Lam; SK01, Sai Kung; SWH, Sheung Wo Hang, Sha Tau Kok; TLC01, Tai Lam-Shek Kong; TLC13, Tai Lam-Shek Kong; WKT, Wu Kau Tang; YSO, Yung Shue O Stream, Sai Kun.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="viruses-11-00423-t002" orientation="portrait" position="float">
<object-id pub-id-type="pii">viruses-11-00423-t002_Table 2</object-id>
<label>Table 2</label>
<caption>
<p>Coding potential and predicted domains in different proteins of Rs-BatCoV HKU32 strain TLC28A.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th colspan="2" align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1">Putative TRS</th>
</tr>
<tr>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">ORF</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nucleotide Positions (Start–End)</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">No. of Nucleotides</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">No. of Amino Acids</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Frame(s)</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nucleotide Position in Genome</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TRS Sequence (Distance (No. of Bases) to AUG)
<sup>1</sup>
</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">1ab</td>
<td align="left" valign="middle" rowspan="1" colspan="1">291–20,428</td>
<td align="left" valign="middle" rowspan="1" colspan="1">20,137</td>
<td align="left" valign="middle" rowspan="1" colspan="1">6712</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2, +3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">69</td>
<td align="left" valign="middle" rowspan="1" colspan="1">AA
<bold>CUAAAC</bold>
(216)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">291–875</td>
<td align="left" valign="middle" rowspan="1" colspan="1">585</td>
<td align="left" valign="middle" rowspan="1" colspan="1">195</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">876–2963</td>
<td align="left" valign="middle" rowspan="1" colspan="1">2088</td>
<td align="left" valign="middle" rowspan="1" colspan="1">696</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">2964–7649</td>
<td align="left" valign="middle" rowspan="1" colspan="1">4686</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1562</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nps4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">7650–9083</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1434</td>
<td align="left" valign="middle" rowspan="1" colspan="1">478</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">9084–9989</td>
<td align="left" valign="middle" rowspan="1" colspan="1">906</td>
<td align="left" valign="middle" rowspan="1" colspan="1">302</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">9990–10817</td>
<td align="left" valign="middle" rowspan="1" colspan="1">828</td>
<td align="left" valign="middle" rowspan="1" colspan="1">276</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp7</td>
<td align="left" valign="middle" rowspan="1" colspan="1">10,818–11,066</td>
<td align="left" valign="middle" rowspan="1" colspan="1">249</td>
<td align="left" valign="middle" rowspan="1" colspan="1">83</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">11,067–11,651</td>
<td align="left" valign="middle" rowspan="1" colspan="1">585</td>
<td align="left" valign="middle" rowspan="1" colspan="1">195</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp9</td>
<td align="left" valign="middle" rowspan="1" colspan="1">11,652–11,975</td>
<td align="left" valign="middle" rowspan="1" colspan="1">324</td>
<td align="left" valign="middle" rowspan="1" colspan="1">108</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp10</td>
<td align="left" valign="middle" rowspan="1" colspan="1">11,976–12,383</td>
<td align="left" valign="middle" rowspan="1" colspan="1">408</td>
<td align="left" valign="middle" rowspan="1" colspan="1">136</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp11</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1">51</td>
<td align="left" valign="middle" rowspan="1" colspan="1">17</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp12</td>
<td align="left" valign="middle" rowspan="1" colspan="1">12,384–15,163</td>
<td align="left" valign="middle" rowspan="1" colspan="1">2780</td>
<td align="left" valign="middle" rowspan="1" colspan="1">927</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp13</td>
<td align="left" valign="middle" rowspan="1" colspan="1">15,164–16,954</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1791</td>
<td align="left" valign="middle" rowspan="1" colspan="1">597</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp14</td>
<td align="left" valign="middle" rowspan="1" colspan="1">16,955–18,508</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1554</td>
<td align="left" valign="middle" rowspan="1" colspan="1">518</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp15</td>
<td align="left" valign="middle" rowspan="1" colspan="1">18,509–19,525</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1017</td>
<td align="left" valign="middle" rowspan="1" colspan="1">339</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp16</td>
<td align="left" valign="middle" rowspan="1" colspan="1">19,526–20,428</td>
<td align="left" valign="middle" rowspan="1" colspan="1">903</td>
<td align="left" valign="middle" rowspan="1" colspan="1">300</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">20,430–24,485</td>
<td align="left" valign="middle" rowspan="1" colspan="1">4056</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1351</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">20,421</td>
<td align="left" valign="middle" rowspan="1" colspan="1">AA
<bold>CUAAAU</bold>
(3)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">ORF3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">24,485–25,153</td>
<td align="left" valign="middle" rowspan="1" colspan="1">669</td>
<td align="left" valign="middle" rowspan="1" colspan="1">222</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">24,279</td>
<td align="left" valign="middle" rowspan="1" colspan="1">TC
<bold>CUUAAC</bold>
(199)
<bold>AUG</bold>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">ORF4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">25,184–25,543</td>
<td align="left" valign="middle" rowspan="1" colspan="1">360</td>
<td align="left" valign="middle" rowspan="1" colspan="1">119</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">ORF5a</td>
<td align="left" valign="middle" rowspan="1" colspan="1">25,544–25,888</td>
<td align="left" valign="middle" rowspan="1" colspan="1">345</td>
<td align="left" valign="middle" rowspan="1" colspan="1">114</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">25,540</td>
<td align="left" valign="middle" rowspan="1" colspan="1">GA
<bold>CUAAAU</bold>
G</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">ORF5b</td>
<td align="left" valign="middle" rowspan="1" colspan="1">25,782–26,225</td>
<td align="left" valign="middle" rowspan="1" colspan="1">444</td>
<td align="left" valign="middle" rowspan="1" colspan="1">147</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">E</td>
<td align="left" valign="middle" rowspan="1" colspan="1">26,209–26,433</td>
<td align="left" valign="middle" rowspan="1" colspan="1">225</td>
<td align="left" valign="middle" rowspan="1" colspan="1">74</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">26,140</td>
<td align="left" valign="middle" rowspan="1" colspan="1">AA
<bold>CUAAAC</bold>
(64)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">M</td>
<td align="left" valign="middle" rowspan="1" colspan="1">26,440–27,126</td>
<td align="left" valign="middle" rowspan="1" colspan="1">687</td>
<td align="left" valign="middle" rowspan="1" colspan="1">228</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">26,430</td>
<td align="left" valign="middle" rowspan="1" colspan="1">GT
<bold>CUAAAC</bold>
(4)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">N</td>
<td align="left" valign="middle" rowspan="1" colspan="1">27,137–28,279</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1143</td>
<td align="left" valign="middle" rowspan="1" colspan="1">380</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">27,128</td>
<td align="left" valign="middle" rowspan="1" colspan="1">AA
<bold>CUAAAC</bold>
(3)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">ORF9</td>
<td align="left" valign="middle" rowspan="1" colspan="1">28,251–28,568</td>
<td align="left" valign="middle" rowspan="1" colspan="1">318</td>
<td align="left" valign="middle" rowspan="1" colspan="1">105</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">28,187</td>
<td align="left" valign="middle" rowspan="1" colspan="1">AG
<bold>CUGAAC</bold>
(58)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">ORF10 (SARS-CoV ORF7a-like protein)</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">28,593–28,955</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">363</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">120</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">28,284</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AA
<bold>CUAAAC</bold>
(303)AUG</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>
<sup>1</sup>
TRS sequences are shown in bold. </p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="viruses-11-00423-t003" orientation="portrait" position="float">
<object-id pub-id-type="pii">viruses-11-00423-t003_Table 3</object-id>
<label>Table 3</label>
<caption>
<p>Characteristics of putative nonstructural proteins of open reading frame (ORF)1ab in Rs-BatCoV HKU32 strain TLC28A, Tr-BatCoV HKU33 strain GZ151867, BatCoV HKU10 and BtNv-AlphaCoV/SC2013.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Amino Acids</th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1"></th>
</tr>
<tr>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">nsp</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Putative Function or Domain</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rs-BatCoV HKU32
<break></break>
Strain TLC28A</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Ro-BatCoV HKU10 183A</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tr-BatCoV HKU33
<break></break>
Strain GZ151867</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">BtNv-AlphaCoV/SC2013</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Unknown</td>
<td align="left" valign="middle" rowspan="1" colspan="1">M
<sup>1</sup>
– A
<sup>195</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">M
<sup>1</sup>
– A
<sup>195</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">M
<sup>1</sup>
– A
<sup>193</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">M
<sup>1</sup>
– A
<sup>193</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Unknown</td>
<td align="left" valign="middle" rowspan="1" colspan="1">P
<sup>196</sup>
– G
<sup>891</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">K
<sup>196</sup>
– G
<sup>888</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">K
<sup>194</sup>
– G
<sup>771</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">K
<sup>194</sup>
– G
<sup>771</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">ADRP, Putative PL
<sup>pro</sup>
<break></break>
domains (PL1
<sup>pro</sup>
, PL2
<sup>pro</sup>
)</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G
<sup>892</sup>
– G
<sup>2453</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>889</sup>
– G
<sup>2518</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G
<sup>772</sup>
– G
<sup>2339</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G
<sup>772</sup>
– G
<sup>2338</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Hydrophobic domain</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>2454</sup>
– Q
<sup>2931</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>2519</sup>
– Q
<sup>2996</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G
<sup>2340</sup>
– Q
<sup>2817</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G
<sup>2339</sup>
– Q
<sup>2815</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">3CL
<sup>pro</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>2932</sup>
– Q
<sup>3233</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>2997</sup>
– Q
<sup>3298</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>2818</sup>
– Q
<sup>3119</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>2816</sup>
– Q
<sup>3117</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Hydrophobic domain</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3234</sup>
– Q
<sup>3509</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3299</sup>
– Q
<sup>3574</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G
<sup>3120</sup>
– Q
<sup>3398</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3118</sup>
– Q
<sup>3395</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp7</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Unknown</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3510</sup>
– Q
<sup>3592</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3575</sup>
– Q
<sup>3657</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3399</sup>
– Q
<sup>3481</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3396</sup>
– Q
<sup>3478</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Unknown</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3593</sup>
– Q
<sup>3787</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3658</sup>
– Q
<sup>3852</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3482</sup>
– Q
<sup>3676</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>3479</sup>
– Q
<sup>3673</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp9</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Unknown</td>
<td align="left" valign="middle" rowspan="1" colspan="1">N
<sup>3788</sup>
– Q
<sup>3895</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">N
<sup>3853</sup>
– Q
<sup>3960</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">N
<sup>3677</sup>
– Q
<sup>3784</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">N
<sup>3674</sup>
– Q
<sup>3781</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp10</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Unknown</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>3896</sup>
– Q
<sup>4031</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>3961</sup>
– Q
<sup>4097</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>3785</sup>
– Q
<sup>3919</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>3782</sup>
–Q
<sup>3916</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp11</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Unknown</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>4032</sup>
– D
<sup>4048</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>4098</sup>
– Q
<sup>4115</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">T
<sup>3920</sup>
– D
<sup>3936</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>3917</sup>
– D
<sup>3933</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp12</td>
<td align="left" valign="middle" rowspan="1" colspan="1">RdRp</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>4032</sup>
– Q
<sup>4958</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>4098</sup>
– Q
<sup>5024</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">T
<sup>3920</sup>
– Q
<sup>4846</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>3917</sup>
– Q
<sup>4843</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp13</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Hel</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>4959</sup>
– Q
<sup>5555</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>5025</sup>
– Q
<sup>5621</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>4847</sup>
– Q
<sup>5443</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>4844</sup>
– Q
<sup>5440</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp14</td>
<td align="left" valign="middle" rowspan="1" colspan="1">ExoN, N7-MTase</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>5556</sup>
– Q
<sup>6073</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>5622</sup>
– Q
<sup>6139</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A
<sup>5444</sup>
– Q
<sup>5960</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>5441</sup>
– Q
<sup>5958</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp15</td>
<td align="left" valign="middle" rowspan="1" colspan="1">NendoU</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G
<sup>6074</sup>
– Q
<sup>6412</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>6140</sup>
– Q
<sup>6478</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">S
<sup>5961</sup>
– Q
<sup>6299</sup>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G
<sup>5959</sup>
– Q
<sup>6297</sup>
</td>
</tr>
<tr>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">nsp16</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">O-MT</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">A
<sup>6413</sup>
– K
<sup>6712</sup>
</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">S
<sup>6479</sup>
– R
<sup>6780</sup>
</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">S
<sup>6300</sup>
– Y
<sup>6591</sup>
</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">S
<sup>6298</sup>
– Y
<sup>6589</sup>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="viruses-11-00423-t004" orientation="portrait" position="float">
<object-id pub-id-type="pii">viruses-11-00423-t004_Table 4</object-id>
<label>Table 4</label>
<caption>
<p>Cleavage site used between nsp in Rs-BatCoV HKU32 strain TLC28A, Tr-BatCoV HKU33 strain GZ151867, BatCoV HKU10 and BtNv-AlphaCoV/SC2013.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th rowspan="2" align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">nsp</th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Cleavage Site</th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1"></th>
</tr>
<tr>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rs-BatCoV HKU32 Strain TLC28A</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Ro-BatCoV HKU10 183A</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tr-BatCoV HKU33 Strain GZ151867</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">BtNv-AlphaCoV/SC2013</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp1/nsp2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A/P</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A/K</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A/K</td>
<td align="left" valign="middle" rowspan="1" colspan="1">A/K</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp2/nsp3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G/G</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G/G</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G/G</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp3/nsp4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G/G</td>
<td align="left" valign="middle" rowspan="1" colspan="1">G/G</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp4/nsp5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/A</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp5/nsp6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/G</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp6/nsp7</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp7/nsp8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp8/nsp9</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/N</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/N</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/N</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/N</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp9/nsp10</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/A</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/A</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/A</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/A</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp10/nsp12</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/A</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/T</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/A</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp12/nsp13</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/A</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp13/nsp14</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/A</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/A</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp14/nsp15</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/G</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Q/G</td>
</tr>
<tr>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">nsp15/nsp16</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Q/A</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Q/S</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Q/S</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="viruses-11-00423-t005" orientation="portrait" position="float">
<object-id pub-id-type="pii">viruses-11-00423-t005_Table 5</object-id>
<label>Table 5</label>
<caption>
<p>Pairwise comparison of
<italic>Coronaviridae</italic>
-wide conserved domains in replicase polyprotein 1ab and overall replicase polyprotein 1ab between Rs-BatCoV HKU32 strain TLC28A and other alphaCoVs.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th rowspan="2" align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Replicase
<break></break>
Polyprotein Domain</th>
<th colspan="5" align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1">Pairwise Sequence Identity with Rs-BatCoV HKU32 Strain TLC28A (%)</th>
</tr>
<tr>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">BtRf-AlphaCoV/
<break></break>
HuB2013</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">BtMs-AlphaCoV/
<break></break>
GS2013</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Ro-BatCoV HKU10</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PEDV</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tr-BatCoV HKU33 strain GZ151867</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">67.6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">67.6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">60.3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">50.1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">49.0</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">84.8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">84.8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">81.5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">74.2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">75.2</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp12</td>
<td align="left" valign="middle" rowspan="1" colspan="1">92.6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">92.6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">90.1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">83.2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">83.7</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp13</td>
<td align="left" valign="middle" rowspan="1" colspan="1">94.1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">94.3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">92.1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">85.6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">80.9</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp14</td>
<td align="left" valign="middle" rowspan="1" colspan="1">93.4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">93.4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">90.0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">79.9</td>
<td align="left" valign="middle" rowspan="1" colspan="1">78.8</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp15</td>
<td align="left" valign="middle" rowspan="1" colspan="1">89.4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">89.4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">83.8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">76.7</td>
<td align="left" valign="middle" rowspan="1" colspan="1">78.2</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp16</td>
<td align="left" valign="middle" rowspan="1" colspan="1">89.7</td>
<td align="left" valign="middle" rowspan="1" colspan="1">90.0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">85.8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">82.8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">81.2</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">7 Concatenated Domains</td>
<td align="left" valign="middle" rowspan="1" colspan="1">83.2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">83.3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">78.6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">70.3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">69.5</td>
</tr>
<tr>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Overall replicase pp1ab</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">80.1</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">80.5</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">75.0</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">65.8</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">63.3</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="viruses-11-00423-t006" orientation="portrait" position="float">
<object-id pub-id-type="pii">viruses-11-00423-t006_Table 6</object-id>
<label>Table 6</label>
<caption>
<p>Coding potential and predicted domains in different proteins of Tr-BatCoV HKU33 strain GZ151867.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th align="left" valign="middle" style="border-top:solid thin" rowspan="1" colspan="1"></th>
<th colspan="2" align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1">Putative TRS</th>
</tr>
<tr>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">ORF</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nucleotide Positions (Start–End)</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">No. of Nucleotides</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">No. of Amino Acids</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Frame(s)</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nucleotide Position in Genome</th>
<th align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TRS Sequence (Distance (No. of Bases) to AUG)
<sup>1</sup>
</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">1ab</td>
<td align="left" valign="middle" rowspan="1" colspan="1">278–20,052</td>
<td align="left" valign="middle" rowspan="1" colspan="1">19,774</td>
<td align="left" valign="middle" rowspan="1" colspan="1">6591</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+1, +2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">54</td>
<td align="left" valign="middle" rowspan="1" colspan="1">AA
<bold>CUAAAC</bold>
(218)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">278–856</td>
<td align="left" valign="middle" rowspan="1" colspan="1">579</td>
<td align="left" valign="middle" rowspan="1" colspan="1">193</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">857–2590</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1734</td>
<td align="left" valign="middle" rowspan="1" colspan="1">578</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">2591–7294</td>
<td align="left" valign="middle" rowspan="1" colspan="1">4704</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1568</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nps4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">7295–8728</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1434</td>
<td align="left" valign="middle" rowspan="1" colspan="1">478</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">8729–9634</td>
<td align="left" valign="middle" rowspan="1" colspan="1">906</td>
<td align="left" valign="middle" rowspan="1" colspan="1">302</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">9635–10,471</td>
<td align="left" valign="middle" rowspan="1" colspan="1">837</td>
<td align="left" valign="middle" rowspan="1" colspan="1">279</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp7</td>
<td align="left" valign="middle" rowspan="1" colspan="1">10,472–10,532</td>
<td align="left" valign="middle" rowspan="1" colspan="1">249</td>
<td align="left" valign="middle" rowspan="1" colspan="1">83</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">10,533–11,305</td>
<td align="left" valign="middle" rowspan="1" colspan="1">585</td>
<td align="left" valign="middle" rowspan="1" colspan="1">195</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp9</td>
<td align="left" valign="middle" rowspan="1" colspan="1">11,306–11,629</td>
<td align="left" valign="middle" rowspan="1" colspan="1">324</td>
<td align="left" valign="middle" rowspan="1" colspan="1">108</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp10</td>
<td align="left" valign="middle" rowspan="1" colspan="1">11,630–12,034</td>
<td align="left" valign="middle" rowspan="1" colspan="1">405</td>
<td align="left" valign="middle" rowspan="1" colspan="1">135</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp11</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1">51</td>
<td align="left" valign="middle" rowspan="1" colspan="1">17</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp12</td>
<td align="left" valign="middle" rowspan="1" colspan="1">12,035–14,814</td>
<td align="left" valign="middle" rowspan="1" colspan="1">2780</td>
<td align="left" valign="middle" rowspan="1" colspan="1">927</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+1</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp13</td>
<td align="left" valign="middle" rowspan="1" colspan="1">14,815–16,605</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1791</td>
<td align="left" valign="middle" rowspan="1" colspan="1">597</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+1</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp14</td>
<td align="left" valign="middle" rowspan="1" colspan="1">16,606–18,156</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1551</td>
<td align="left" valign="middle" rowspan="1" colspan="1">517</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+1</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp15</td>
<td align="left" valign="middle" rowspan="1" colspan="1">18,157–19,173</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1017</td>
<td align="left" valign="middle" rowspan="1" colspan="1">339</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+1</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp16</td>
<td align="left" valign="middle" rowspan="1" colspan="1">19,174–20,052</td>
<td align="left" valign="middle" rowspan="1" colspan="1">879</td>
<td align="left" valign="middle" rowspan="1" colspan="1">292</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+1</td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
<td align="left" valign="middle" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">S</td>
<td align="left" valign="middle" rowspan="1" colspan="1">20,053–24,150</td>
<td align="left" valign="middle" rowspan="1" colspan="1">4098</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1365</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">20,049</td>
<td align="left" valign="middle" rowspan="1" colspan="1">GA
<bold>CUAAAU</bold>
G</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">ORF3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">24,150–24,755</td>
<td align="left" valign="middle" rowspan="1" colspan="1">606</td>
<td align="left" valign="middle" rowspan="1" colspan="1">201</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">23,876</td>
<td align="left" valign="middle" rowspan="1" colspan="1">AT
<bold>CUCAAC</bold>
(268)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">E</td>
<td align="left" valign="middle" rowspan="1" colspan="1">24,777–25,004</td>
<td align="left" valign="middle" rowspan="1" colspan="1">228</td>
<td align="left" valign="middle" rowspan="1" colspan="1">75</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">24,763</td>
<td align="left" valign="middle" rowspan="1" colspan="1">TT
<bold>CUCAAC</bold>
(8)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">M</td>
<td align="left" valign="middle" rowspan="1" colspan="1">25,011–25,697</td>
<td align="left" valign="middle" rowspan="1" colspan="1">687</td>
<td align="left" valign="middle" rowspan="1" colspan="1">228</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">25,001</td>
<td align="left" valign="middle" rowspan="1" colspan="1">GT
<bold>CUAAAC</bold>
(4)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">N</td>
<td align="left" valign="middle" rowspan="1" colspan="1">25,706–26,977</td>
<td align="left" valign="middle" rowspan="1" colspan="1">1272</td>
<td align="left" valign="middle" rowspan="1" colspan="1">423</td>
<td align="left" valign="middle" rowspan="1" colspan="1">+2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">25,699</td>
<td align="left" valign="middle" rowspan="1" colspan="1">AA
<bold>CUAAAC</bold>
(1)AUG</td>
</tr>
<tr>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">ORF7</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">26,989–27,348</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">360</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">119</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">+1</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">26,982</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AA
<bold>CUAAAU</bold>
(1)AUG</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>
<sup>1</sup>
TRS sequences are shown in bold.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="viruses-11-00423-t007" orientation="portrait" position="float">
<object-id pub-id-type="pii">viruses-11-00423-t007_Table 7</object-id>
<label>Table 7</label>
<caption>
<p>Pairwise comparison of
<italic>Coronaviridae</italic>
-wide conserved domains in replicase polyprotein 1ab and overall replicase polyprotein 1ab between Tr-BatCoV HKU33 strain GZ151867 and other alphaCoVs.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th rowspan="2" align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Replicase Polyprotein Domain</th>
<th colspan="5" align="left" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1">Pairwise Amino Acid Sequence Identity with Tr-BatCoV HKU33 Strain GZ151867 (%)</th>
</tr>
<tr>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">BtNv-AlphaCoV/SC2013</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">BtRf-AlphaCoV/HuB2013</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">BtMs-AlphaCoV/GS2013</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AlphaCoV BatCoV/P.kuhlii/Italy
<break></break>
206679-3/2010</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rs-BatCoV HKU32 Strain TLC28A</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">58.8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">48.7</td>
<td align="left" valign="middle" rowspan="1" colspan="1">48.8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">57.3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">49.0</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">82.5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">74.2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">74.5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">78.8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">75.2</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp12</td>
<td align="left" valign="middle" rowspan="1" colspan="1">86.3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">83.5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">83.6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">86.4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">83.7</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp13</td>
<td align="left" valign="middle" rowspan="1" colspan="1">87.0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">79.7</td>
<td align="left" valign="middle" rowspan="1" colspan="1">80.1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">82.7</td>
<td align="left" valign="middle" rowspan="1" colspan="1">80.9</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp14</td>
<td align="left" valign="middle" rowspan="1" colspan="1">84.0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">79.3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">79.3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">82.4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">78.8</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp15</td>
<td align="left" valign="middle" rowspan="1" colspan="1">85.8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">79.6</td>
<td align="left" valign="middle" rowspan="1" colspan="1">79.0</td>
<td align="left" valign="middle" rowspan="1" colspan="1">84.7</td>
<td align="left" valign="middle" rowspan="1" colspan="1">78.2</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">nsp16</td>
<td align="left" valign="middle" rowspan="1" colspan="1">87.3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">80.8</td>
<td align="left" valign="middle" rowspan="1" colspan="1">80.5</td>
<td align="left" valign="middle" rowspan="1" colspan="1">84.2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">81.2</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="1" colspan="1">7 Concatenated Domains</td>
<td align="left" valign="middle" rowspan="1" colspan="1">76.3</td>
<td align="left" valign="middle" rowspan="1" colspan="1">69.2</td>
<td align="left" valign="middle" rowspan="1" colspan="1">69.1</td>
<td align="left" valign="middle" rowspan="1" colspan="1">74.4</td>
<td align="left" valign="middle" rowspan="1" colspan="1">69.4</td>
</tr>
<tr>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Overall replicase pp1ab</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">73.4</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">62.9</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">63.0</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">71.6</td>
<td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">63.3</td>
</tr>
</tbody>
</table>
</table-wrap>
</floats-group>
</pmc>
</record>

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