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Bat Coronaviruses in China

Identifieur interne : 000488 ( Pmc/Checkpoint ); précédent : 000487; suivant : 000489

Bat Coronaviruses in China

Auteurs : Yi Fan [République populaire de Chine] ; Kai Zhao [République populaire de Chine] ; Zheng-Li Shi [République populaire de Chine] ; Peng Zhou [République populaire de Chine]

Source :

RBID : PMC:6466186

Abstract

During the past two decades, three zoonotic coronaviruses have been identified as the cause of large-scale disease outbreaks–Severe Acute Respiratory Syndrome (SARS), Middle East Respiratory Syndrome (MERS), and Swine Acute Diarrhea Syndrome (SADS). SARS and MERS emerged in 2003 and 2012, respectively, and caused a worldwide pandemic that claimed thousands of human lives, while SADS struck the swine industry in 2017. They have common characteristics, such as they are all highly pathogenic to humans or livestock, their agents originated from bats, and two of them originated in China. Thus, it is highly likely that future SARS- or MERS-like coronavirus outbreaks will originate from bats, and there is an increased probability that this will occur in China. Therefore, the investigation of bat coronaviruses becomes an urgent issue for the detection of early warning signs, which in turn minimizes the impact of such future outbreaks in China. The purpose of the review is to summarize the current knowledge on viral diversity, reservoir hosts, and the geographical distributions of bat coronaviruses in China, and eventually we aim to predict virus hotspots and their cross-species transmission potential.


Url:
DOI: 10.3390/v11030210
PubMed: 30832341
PubMed Central: 6466186


Affiliations:


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PMC:6466186

Le document en format XML

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<p>During the past two decades, three zoonotic coronaviruses have been identified as the cause of large-scale disease outbreaks–Severe Acute Respiratory Syndrome (SARS), Middle East Respiratory Syndrome (MERS), and Swine Acute Diarrhea Syndrome (SADS). SARS and MERS emerged in 2003 and 2012, respectively, and caused a worldwide pandemic that claimed thousands of human lives, while SADS struck the swine industry in 2017. They have common characteristics, such as they are all highly pathogenic to humans or livestock, their agents originated from bats, and two of them originated in China. Thus, it is highly likely that future SARS- or MERS-like coronavirus outbreaks will originate from bats, and there is an increased probability that this will occur in China. Therefore, the investigation of bat coronaviruses becomes an urgent issue for the detection of early warning signs, which in turn minimizes the impact of such future outbreaks in China. The purpose of the review is to summarize the current knowledge on viral diversity, reservoir hosts, and the geographical distributions of bat coronaviruses in China, and eventually we aim to predict virus hotspots and their cross-species transmission potential.</p>
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</TEI>
<pmc article-type="review-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">30832341</article-id>
<article-id pub-id-type="pmc">6466186</article-id>
<article-id pub-id-type="doi">10.3390/v11030210</article-id>
<article-id pub-id-type="publisher-id">viruses-11-00210</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Review</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Bat Coronaviruses in China</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0002-4117-8433</contrib-id>
<name>
<surname>Fan</surname>
<given-names>Yi</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00210">1</xref>
<xref ref-type="aff" rid="af2-viruses-11-00210">2</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zhao</surname>
<given-names>Kai</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00210">1</xref>
<xref ref-type="aff" rid="af2-viruses-11-00210">2</xref>
</contrib>
<contrib contrib-type="author">
<contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0001-8089-163X</contrib-id>
<name>
<surname>Shi</surname>
<given-names>Zheng-Li</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00210">1</xref>
<xref ref-type="aff" rid="af2-viruses-11-00210">2</xref>
</contrib>
<contrib contrib-type="author">
<contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0001-9863-4201</contrib-id>
<name>
<surname>Zhou</surname>
<given-names>Peng</given-names>
</name>
<xref ref-type="aff" rid="af1-viruses-11-00210">1</xref>
<xref ref-type="aff" rid="af2-viruses-11-00210">2</xref>
<xref rid="c1-viruses-11-00210" ref-type="corresp">*</xref>
</contrib>
</contrib-group>
<aff id="af1-viruses-11-00210">
<label>1</label>
CAS Key Laboratory of Special Pathogens and Biosafety, Wuhan Institute of Virology, Chinese Academy of Sciences, Wuhan 430071, China;
<email>yifanfs0224@163.com</email>
(Y.F.);
<email>chiukal@163.com</email>
(K.Z.);
<email>zlshi@wh.iov.cn</email>
(Z.-L.S.)</aff>
<aff id="af2-viruses-11-00210">
<label>2</label>
University of Chinese Academy of Sciences, Beijing 100049, China</aff>
<author-notes>
<corresp id="c1-viruses-11-00210">
<label>*</label>
Correspondence:
<email>peng.zhou@wh.iov.cn</email>
</corresp>
</author-notes>
<pub-date pub-type="epub">
<day>02</day>
<month>3</month>
<year>2019</year>
</pub-date>
<pub-date pub-type="collection">
<month>3</month>
<year>2019</year>
</pub-date>
<volume>11</volume>
<issue>3</issue>
<elocation-id>210</elocation-id>
<history>
<date date-type="received">
<day>29</day>
<month>1</month>
<year>2019</year>
</date>
<date date-type="accepted">
<day>26</day>
<month>2</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>During the past two decades, three zoonotic coronaviruses have been identified as the cause of large-scale disease outbreaks–Severe Acute Respiratory Syndrome (SARS), Middle East Respiratory Syndrome (MERS), and Swine Acute Diarrhea Syndrome (SADS). SARS and MERS emerged in 2003 and 2012, respectively, and caused a worldwide pandemic that claimed thousands of human lives, while SADS struck the swine industry in 2017. They have common characteristics, such as they are all highly pathogenic to humans or livestock, their agents originated from bats, and two of them originated in China. Thus, it is highly likely that future SARS- or MERS-like coronavirus outbreaks will originate from bats, and there is an increased probability that this will occur in China. Therefore, the investigation of bat coronaviruses becomes an urgent issue for the detection of early warning signs, which in turn minimizes the impact of such future outbreaks in China. The purpose of the review is to summarize the current knowledge on viral diversity, reservoir hosts, and the geographical distributions of bat coronaviruses in China, and eventually we aim to predict virus hotspots and their cross-species transmission potential.</p>
</abstract>
<kwd-group>
<kwd>coronavirus</kwd>
<kwd>bat</kwd>
<kwd>epidemiology</kwd>
<kwd>cross-species</kwd>
<kwd>zoonosis</kwd>
</kwd-group>
</article-meta>
</front>
<floats-group>
<fig id="viruses-11-00210-f001" orientation="portrait" position="float">
<label>Figure 1</label>
<caption>
<p>Geographical distribution of bat coronaviruses (CoVs) and their corresponding bat hosts in China. Each red box represents one CoV positive sample found in that particular bat species. One dot matrix was drawn for each province where a CoV positive sample had been reported. Guangdong Province, where SARS and SADS began, is circled in red. Abbreviations of bat species and virus species are indicated.</p>
</caption>
<graphic xlink:href="viruses-11-00210-g001"></graphic>
</fig>
<fig id="viruses-11-00210-f002" orientation="portrait" position="float">
<label>Figure 2</label>
<caption>
<p>Genetic diversity of bat CoV in China. Sequences of 440 bp conserved the viral polymerase (RdRp) region for each CoV species were compared to related reference sequences. Reference genomes used: BtCoV-HKU10, NC_018871.1; BtRfCoV-HuB13, NC_028814.1; BtMiCoV-1, EU420138.1; BtMiCoV-HKU8, NC_010438.1; BtRhCoV-HKU2, MF094682.1; BtHpCoV-ZJ13, NC_025217.1; MERSr-CoV, NC_038294.1; BtPiCoV-HKU5, NC_009020.1; BtTyCoV-HKU4, NC_009019.1; BtRoCoV-GCCDC1, MG762606.1; BtRoCoV-HKU9, NC_009021.1; and SARSr-CoV, NC_004718.3. Notably, samples that were positive for BtMy-Sax11, BtNy-Sc13, and BtScCoV-512 were also identified in China. These were not taken into account here as too few sequences were available.</p>
</caption>
<graphic xlink:href="viruses-11-00210-g002"></graphic>
</fig>
<table-wrap id="viruses-11-00210-t001" orientation="portrait" position="float">
<object-id pub-id-type="pii">viruses-11-00210-t001_Table 1</object-id>
<label>Table 1</label>
<caption>
<p>International Committee of Taxonomy of Viruses (ICTV) classification of coronaviruses species, reservoir hosts, and presence reported in China.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Coronavirus Species</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Abbreviations</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Human</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Bats</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Other Animals</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Reported in China</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1"></th>
</tr>
</thead>
<tbody>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Bat coronavirus HKU10</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtCoV-HKU10</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B7-viruses-11-00210" ref-type="bibr">7</xref>
,
<xref rid="B8-viruses-11-00210" ref-type="bibr">8</xref>
,
<xref rid="B26-viruses-11-00210" ref-type="bibr">26</xref>
,
<xref rid="B27-viruses-11-00210" ref-type="bibr">27</xref>
]</td>
<td rowspan="17" align="center" valign="middle" style="border-bottom:solid thin" colspan="1">
<bold>α-CoV</bold>
</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Bat coronavirus CDPHE15</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtCoV-CDPHE15</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">No</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<italic>Rhinolophus ferrumequinum</italic>
alphacoronavirus HuB-2013</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtRfCoV-HuB13</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B8-viruses-11-00210" ref-type="bibr">8</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">* Human coronavirus 229E</td>
<td align="center" valign="middle" rowspan="1" colspan="1">HCoV-229E</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B28-viruses-11-00210" ref-type="bibr">28</xref>
,
<xref rid="B29-viruses-11-00210" ref-type="bibr">29</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Lucheng Rn rat coronavirus</td>
<td align="center" valign="middle" rowspan="1" colspan="1">LRNV</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (rat)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B30-viruses-11-00210" ref-type="bibr">30</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Ferret coronavirus</td>
<td align="center" valign="middle" rowspan="1" colspan="1">FRCoV</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (ferret)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">No [
<xref rid="B31-viruses-11-00210" ref-type="bibr">31</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">* Mink coronavirus 1</td>
<td align="center" valign="middle" rowspan="1" colspan="1">MCoV</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (mink)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">No [
<xref rid="B14-viruses-11-00210" ref-type="bibr">14</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<italic>Miniopterus</italic>
bat coronavirus 1</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtMiCoV-1</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B7-viruses-11-00210" ref-type="bibr">7</xref>
,
<xref rid="B8-viruses-11-00210" ref-type="bibr">8</xref>
,
<xref rid="B32-viruses-11-00210" ref-type="bibr">32</xref>
,
<xref rid="B33-viruses-11-00210" ref-type="bibr">33</xref>
,
<xref rid="B34-viruses-11-00210" ref-type="bibr">34</xref>
,
<xref rid="B35-viruses-11-00210" ref-type="bibr">35</xref>
,
<xref rid="B36-viruses-11-00210" ref-type="bibr">36</xref>
,
<xref rid="B37-viruses-11-00210" ref-type="bibr">37</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<italic>Miniopterus</italic>
bat coronavirus HKU8</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtMiCoV-HKU8</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B7-viruses-11-00210" ref-type="bibr">7</xref>
,
<xref rid="B8-viruses-11-00210" ref-type="bibr">8</xref>
,
<xref rid="B33-viruses-11-00210" ref-type="bibr">33</xref>
,
<xref rid="B34-viruses-11-00210" ref-type="bibr">34</xref>
,
<xref rid="B35-viruses-11-00210" ref-type="bibr">35</xref>
,
<xref rid="B37-viruses-11-00210" ref-type="bibr">37</xref>
,
<xref rid="B38-viruses-11-00210" ref-type="bibr">38</xref>
,
<xref rid="B39-viruses-11-00210" ref-type="bibr">39</xref>
,
<xref rid="B40-viruses-11-00210" ref-type="bibr">40</xref>
,
<xref rid="B41-viruses-11-00210" ref-type="bibr">41</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<italic>Myotis ricketti</italic>
alphacoronavirus Sax-2011</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtMy-Sax11</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B8-viruses-11-00210" ref-type="bibr">8</xref>
,
<xref rid="B37-viruses-11-00210" ref-type="bibr">37</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<italic>Nyctalus velutinus</italic>
alphacoronavirus SC-2013</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtNy-Sc13</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B8-viruses-11-00210" ref-type="bibr">8</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">* Porcine epidemic diarrhea virus</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PEDV</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (pig)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B42-viruses-11-00210" ref-type="bibr">42</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<italic>Scotophilus</italic>
bat coronavirus 512</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtScCoV-512</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B37-viruses-11-00210" ref-type="bibr">37</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<bold>*
<italic>Rhinolophus</italic>
bat coronavirus HKU2 (SADS)</bold>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtRhCoV-HKU2</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B2-viruses-11-00210" ref-type="bibr">2</xref>
,
<xref rid="B7-viruses-11-00210" ref-type="bibr">7</xref>
,
<xref rid="B8-viruses-11-00210" ref-type="bibr">8</xref>
,
<xref rid="B38-viruses-11-00210" ref-type="bibr">38</xref>
,
<xref rid="B43-viruses-11-00210" ref-type="bibr">43</xref>
,
<xref rid="B44-viruses-11-00210" ref-type="bibr">44</xref>
,
<xref rid="B45-viruses-11-00210" ref-type="bibr">45</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">* Human coronavirus NL63</td>
<td align="center" valign="middle" rowspan="1" colspan="1">HCoV-NL63</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B28-viruses-11-00210" ref-type="bibr">28</xref>
,
<xref rid="B29-viruses-11-00210" ref-type="bibr">29</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">NL63-related bat coronavirus strain BtKYNL63-9b</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtKYNL63</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">No [
<xref rid="B24-viruses-11-00210" ref-type="bibr">24</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">* Alphacoronavirus 1 (Transmissible gastroenteritis virus)</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TGEV</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yes (pig)</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yes [
<xref rid="B42-viruses-11-00210" ref-type="bibr">42</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">China
<italic>Rattus</italic>
coronavirus HKU24</td>
<td align="center" valign="middle" rowspan="1" colspan="1">RtCoV-HKU24</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (rat)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B46-viruses-11-00210" ref-type="bibr">46</xref>
]</td>
<td rowspan="12" align="center" valign="middle" style="border-bottom:solid thin" colspan="1">
<bold>β-CoV</bold>
</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">* Human coronavirus HKU1</td>
<td align="center" valign="middle" rowspan="1" colspan="1">HCoV-HKU1</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B28-viruses-11-00210" ref-type="bibr">28</xref>
,
<xref rid="B29-viruses-11-00210" ref-type="bibr">29</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">* Murine coronavirus (Murine hepatitis coronavirus)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">MHV</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (mouse)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">No [
<xref rid="B47-viruses-11-00210" ref-type="bibr">47</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Bat
<italic>Hp</italic>
-betacoronavirus Zhejiang2013</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtHpCoV-ZJ13</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B8-viruses-11-00210" ref-type="bibr">8</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Hedgehog coronavirus 1</td>
<td align="center" valign="middle" rowspan="1" colspan="1">EriCoV-1</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (hedgehog)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">No [
<xref rid="B48-viruses-11-00210" ref-type="bibr">48</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<bold>* Middle East respiratory syndrome-related coronavirus</bold>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">MERSr-CoV</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B49-viruses-11-00210" ref-type="bibr">49</xref>
,
<xref rid="B50-viruses-11-00210" ref-type="bibr">50</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<italic>Pipistrellus</italic>
bat coronavirus HKU5</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtPiCoV-HKU5</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B38-viruses-11-00210" ref-type="bibr">38</xref>
,
<xref rid="B39-viruses-11-00210" ref-type="bibr">39</xref>
,
<xref rid="B49-viruses-11-00210" ref-type="bibr">49</xref>
,
<xref rid="B51-viruses-11-00210" ref-type="bibr">51</xref>
,
<xref rid="B52-viruses-11-00210" ref-type="bibr">52</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<italic>Tylonycteris</italic>
bat coronavirus HKU4</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtTyCoV-HKU4</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B36-viruses-11-00210" ref-type="bibr">36</xref>
,
<xref rid="B38-viruses-11-00210" ref-type="bibr">38</xref>
,
<xref rid="B39-viruses-11-00210" ref-type="bibr">39</xref>
,
<xref rid="B49-viruses-11-00210" ref-type="bibr">49</xref>
,
<xref rid="B50-viruses-11-00210" ref-type="bibr">50</xref>
,
<xref rid="B51-viruses-11-00210" ref-type="bibr">51</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<italic>Rousettus</italic>
bat coronavirus GCCDC1</td>
<td align="center" valign="middle" rowspan="1" colspan="1">
<sup>#</sup>
BtEoCoV-GCCDC1</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B53-viruses-11-00210" ref-type="bibr">53</xref>
,
<xref rid="B54-viruses-11-00210" ref-type="bibr">54</xref>
,
<xref rid="B55-viruses-11-00210" ref-type="bibr">55</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<italic>Rousettus</italic>
bat coronavirus HKU9</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BtRoCoV-HKU9</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B39-viruses-11-00210" ref-type="bibr">39</xref>
,
<xref rid="B55-viruses-11-00210" ref-type="bibr">55</xref>
,
<xref rid="B56-viruses-11-00210" ref-type="bibr">56</xref>
,
<xref rid="B57-viruses-11-00210" ref-type="bibr">57</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">
<bold>* Severe acute respiratory syndrome-related coronavirus</bold>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">SARSr-CoV</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B7-viruses-11-00210" ref-type="bibr">7</xref>
,
<xref rid="B8-viruses-11-00210" ref-type="bibr">8</xref>
,
<xref rid="B20-viruses-11-00210" ref-type="bibr">20</xref>
,
<xref rid="B21-viruses-11-00210" ref-type="bibr">21</xref>
,
<xref rid="B22-viruses-11-00210" ref-type="bibr">22</xref>
,
<xref rid="B27-viruses-11-00210" ref-type="bibr">27</xref>
,
<xref rid="B37-viruses-11-00210" ref-type="bibr">37</xref>
,
<xref rid="B40-viruses-11-00210" ref-type="bibr">40</xref>
,
<xref rid="B45-viruses-11-00210" ref-type="bibr">45</xref>
,
<xref rid="B58-viruses-11-00210" ref-type="bibr">58</xref>
,
<xref rid="B59-viruses-11-00210" ref-type="bibr">59</xref>
,
<xref rid="B60-viruses-11-00210" ref-type="bibr">60</xref>
,
<xref rid="B61-viruses-11-00210" ref-type="bibr">61</xref>
,
<xref rid="B62-viruses-11-00210" ref-type="bibr">62</xref>
,
<xref rid="B63-viruses-11-00210" ref-type="bibr">63</xref>
,
<xref rid="B64-viruses-11-00210" ref-type="bibr">64</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">* Betacoronavirus 1 (Human coronavirus OC43)</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">HCoV-OC43</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yes</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yes [
<xref rid="B28-viruses-11-00210" ref-type="bibr">28</xref>
,
<xref rid="B29-viruses-11-00210" ref-type="bibr">29</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Wigeon coronavirus HKU20</td>
<td align="center" valign="middle" rowspan="1" colspan="1">WiCoV-HKU20</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (bird)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B65-viruses-11-00210" ref-type="bibr">65</xref>
]</td>
<td rowspan="7" align="center" valign="middle" style="border-bottom:solid thin" colspan="1">
<bold>δ-CoV</bold>
</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Bulbul coronavirus HKU11</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BuCoV-HKU11</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (bird)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B65-viruses-11-00210" ref-type="bibr">65</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Coronavirus HKU15</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PoCoV-HKU15</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (pig)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B66-viruses-11-00210" ref-type="bibr">66</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Munia coronavirus HKU13</td>
<td align="center" valign="middle" rowspan="1" colspan="1">MuCoV-HKU13</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (bird)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B65-viruses-11-00210" ref-type="bibr">65</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">White-eye coronavirus HKU16</td>
<td align="center" valign="middle" rowspan="1" colspan="1">WECoV-HKU13</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (bird)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B65-viruses-11-00210" ref-type="bibr">65</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">Night heron coronavirus HKU19</td>
<td align="center" valign="middle" rowspan="1" colspan="1">NHCoV-HKU19</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (bird)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes [
<xref rid="B65-viruses-11-00210" ref-type="bibr">65</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Common moorhen coronavirus HKU21</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CMCoV-HKU21</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yes (bird)</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yes [
<xref rid="B65-viruses-11-00210" ref-type="bibr">65</xref>
]</td>
</tr>
<tr>
<td align="center" valign="middle" rowspan="1" colspan="1">*
<sup>?</sup>
Beluga whale coronavirus SW1</td>
<td align="center" valign="middle" rowspan="1" colspan="1">BWCoV-SW1</td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" rowspan="1" colspan="1">Yes (whale)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">No [
<xref rid="B13-viruses-11-00210" ref-type="bibr">13</xref>
]</td>
<td rowspan="2" align="center" valign="middle" style="border-bottom:solid thin" colspan="1">
<bold>γ-CoV</bold>
</td>
</tr>
<tr>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">* Avian infectious bronchitis virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">IBV</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yes (bird)</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yes [
<xref rid="B12-viruses-11-00210" ref-type="bibr">12</xref>
]</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>* The disease-causing CoVs are indicated and the three zoonotic CoVs are in bold. *? BWCoV-SW1 was found in a sick whale, but whether it was the etiological agent of the infection was not proven. # Carrier of this virus was confirmed as
<italic>Eonycteris spelaea</italic>
, but not
<italic>Rousettus</italic>
bats. The virus was renamed accordingly.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</floats-group>
</pmc>
<affiliations>
<list>
<country>
<li>République populaire de Chine</li>
</country>
<settlement>
<li>Pékin</li>
</settlement>
</list>
<tree>
<country name="République populaire de Chine">
<noRegion>
<name sortKey="Fan, Yi" sort="Fan, Yi" uniqKey="Fan Y" first="Yi" last="Fan">Yi Fan</name>
</noRegion>
<name sortKey="Shi, Zheng Li" sort="Shi, Zheng Li" uniqKey="Shi Z" first="Zheng-Li" last="Shi">Zheng-Li Shi</name>
<name sortKey="Zhao, Kai" sort="Zhao, Kai" uniqKey="Zhao K" first="Kai" last="Zhao">Kai Zhao</name>
<name sortKey="Zhou, Peng" sort="Zhou, Peng" uniqKey="Zhou P" first="Peng" last="Zhou">Peng Zhou</name>
</country>
</tree>
</affiliations>
</record>

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