59:
1902:
31:
669:
The presence of HE genes exclusively in members of
Betacoronavirus subgroup A, but not members of Betacoronavirus subgroup B, C and D suggested that the recombination had probably occurred in the ancestor of members of Betacoronavirus subgroup A, after diverging from the ancestor of other subgroups
561:
CoVs are classified into four genera, Alphacoronavirus, Betacoronavirus, Gammacoronavirus and
Deltacoronavirus. Within Betacoronavirus, they can be further subclassified into lineages A, B, C and D . In 2018, these four lineages were reclassified as subgenera of Betacoronavirus, and renamed as
438:
In all members of
Betacoronavirus subgroup A, a haemagglutinin esterase (HE) gene, which encodes a glycoprotein with neuraminate O-acetyl-esterase activity and the active site FGDS, is present downstream to ORF1ab and upstream to S gene (Figure
562:
Embecovirus (previous lineage A), Sarbecovirus (previous lineage B), Merbecovirus (previous lineage C) and
Nobecovirus (previous lineage D) . In addition, a fifth subgenus, Hibecovirus, was also included (Figure 1) .
684:
Diversity of
Dromedary Camel Coronavirus HKU23 in African Camels Revealed Multiple Recombination Events among Closely Related Betacoronaviruses of the Subgenus Embecovirus. So RTY, et al. J Virol. 2019. PMID
574:
Lai, Michael M. C.; Cavanagh, David (1997-01-01), Maramorosch, Karl; Murphy, Frederick A.; Shatkin, Aaron J. (eds.), "The
Molecular Biology of Coronaviruses; III. Structure of Virions; A. Virion Morphology",
451:
Woo, Patrick C. Y.; Wang, Ming; Lau, Susanna K. P.; Xu, Huifang; Poon, Rosana W. S.; Guo, Rongtong; Wong, Beatrice H. L.; Gao, Kai; Tsoi, Hoi-wah; Huang, Yi; Li, Kenneth S. M. (February 2007).
305:
population. Contributing to this diversity are several recombination events that had taken place in the past between closely related Beta-CoVs of the subgenus
Embecovirus.
278:
structural proteins are anchored. Unlike other coronaviruses, viruses in this subgenus also have an additional shorter spike-like structural protein called
1405:
854:
1511:
1142:
1896:
1980:
1033:
1528:
453:"Comparative Analysis of Twelve Genomes of Three Novel Group 2c and Group 2d Coronaviruses Reveals Unique Group and Subgroup Features"
712:
1905:
1040:
1247:
1431:
1424:
1985:
1756:
1468:
975:
363:
1077:
1070:
1533:
1475:
1312:
191:
968:
1632:
1461:
1242:
271:
267:
1972:
1967:
1777:
705:
58:
2013:
1416:
275:
2008:
1749:
1505:
1340:
212:
1770:
1920:
1669:
1521:
1411:
1398:
1319:
1304:
1135:
698:
279:
247:
198:
1958:
1676:
847:
290:
1901:
53:
8:
45:
1326:
1299:
1237:
1232:
784:
655:
620:
597:
547:
512:
485:
452:
424:
389:
205:
38:
588:
1763:
1662:
1293:
660:
642:
602:
552:
534:
490:
472:
429:
411:
184:
1696:
1579:
794:
650:
632:
592:
584:
542:
524:
511:
Wong, Antonio C. P.; Li, Xin; Lau, Susanna K. P.; Woo, Patrick C. Y. (2019-02-20).
480:
464:
419:
401:
1943:
1332:
1268:
765:
753:
619:
Woo, Patrick C. Y.; Huang, Yi; Lau, Susanna K. P.; Yuen, Kwok-Yung (2010-08-24).
388:
Woo, Patrick C. Y.; Huang, Yi; Lau, Susanna K. P.; Yuen, Kwok-Yung (2010-08-24).
238:
155:
119:
690:
263:
2002:
1590:
954:
870:
741:
723:
646:
538:
476:
415:
302:
262:
The viruses of this subgenus, like other coronaviruses, have a lipid bilayer
143:
95:
1793:
1735:
1491:
1384:
1121:
1093:
1019:
991:
898:
747:
664:
556:
494:
433:
323:
315:
107:
606:
301:
camel beta-coronavirus (Beta-CoV HKU23) exhibits genetic diversity in the
1952:
1834:
1707:
1618:
1447:
1356:
1214:
1186:
1158:
1056:
926:
833:
805:
759:
468:
331:
243:
233:
131:
1648:
1558:
1516:
529:
735:
637:
406:
298:
83:
30:
1914:
1937:
1538:
229:
250:(HE) gene. The viruses in the subgenus were previously known as
1553:
1548:
1543:
355:
294:
42:
70:
360:
International
Committee on Taxonomy of Viruses (ICTV)
1406:
Middle East respiratory syndrome-related coronavirus
855:
Rhinolophus ferrumequinum alphacoronavirus HuB-2013
621:"Coronavirus Genomics and Bioinformatics Analysis"
390:"Coronavirus Genomics and Bioinformatics Analysis"
720:
2000:
1143:NL63-related bat coronavirus strain BtKYNL63-9b
618:
387:
1846:
706:
242:. The viruses in this subgenus, unlike other
510:
450:
1034:Nyctalus velutinus alphacoronavirus SC-2013
573:
1855:
713:
699:
513:"Global Epidemiology of Bat Coronaviruses"
29:
1005:Myotis ricketti alphacoronavirus Sax-2011
654:
636:
596:
546:
528:
484:
423:
405:
680:
678:
297:are present in the same host cell. The
2001:
782:
350:
348:
1919:
1918:
1831:
781:
694:
675:
1041:Pipistrellus kuhlii coronavirus 3398
383:
381:
1370:Bat Hp-betacoronavirus Zhejiang2013
1248:Transmissible gastroenteritis virus
345:
48:, schematic structure, and genome.
13:
612:
14:
2025:
1432:Tylonycteris bat coronavirus HKU4
1425:Pipistrellus bat coronavirus HKU5
567:
378:
366:from the original on 4 March 2018
293:can occur when two or more viral
1900:
1757:Common moorhen coronavirus HKU21
1469:Rousettus bat coronavirus GCCDC1
1107:Rhinolophus bat coronavirus HKU2
976:Miniopterus bat coronavirus HKU8
285:
57:
1832:
1078:Scotophilus bat coronavirus 512
1071:Porcine epidemic diarrhea virus
1534:Bat SARS-like coronavirus WIV1
1476:Rousettus bat coronavirus HKU9
1313:China Rattus coronavirus HKU24
1200:Suncus murinus coronavirus X74
504:
444:
356:"Virus Taxonomy: 2018 Release"
192:China Rattus coronavirus HKU24
1:
1807:Night heron coronavirus HKU19
1172:Sorex araneus coronavirus T14
969:Miniopterus bat coronavirus 1
589:10.1016/S0065-3527(08)60286-9
339:
1633:Beluga whale coronavirus SW1
1462:Eidolon bat coronavirus C704
1243:Canine coronavirus HuPn-2018
257:
7:
1778:White-eye coronavirus HKU16
308:
10:
2030:
912:Lucheng Rn rat coronavirus
577:Advances in Virus Research
1927:
1893:
1864:
1842:
1827:
1802:
1791:
1744:
1733:
1716:
1705:
1694:
1657:
1646:
1627:
1616:
1599:
1588:
1577:
1500:
1489:
1456:
1445:
1417:MERS coronavirus EMC/2012
1393:
1382:
1365:
1354:
1288:
1277:
1266:
1223:
1212:
1195:
1184:
1167:
1156:
1130:
1119:
1102:
1091:
1065:
1054:
1028:
1017:
1000:
989:
963:
952:
935:
924:
907:
896:
879:
868:
842:
831:
814:
803:
792:
777:
731:
180:
175:
52:
37:
28:
21:
1750:Bulbul coronavirus HKU11
1721:Wigeon coronavirus HKU20
1506:SARS-related coronavirus
1341:Myodes coronavirus 2JL14
213:Myodes coronavirus 2JL14
1771:Munia coronavirus HKU13
819:Bat coronavirus CDPHE15
583:, Academic Press: 5–6,
1670:Avian coronavirus 9203
1604:Goose coronavirus CB17
1412:London1_novel CoV/2012
1399:Hedgehog coronavirus 1
1320:Human coronavirus HKU1
1305:Human coronavirus OC43
1136:Human coronavirus NL63
884:Human coronavirus 229E
280:hemagglutinin esterase
252:group 2a coronaviruses
248:hemagglutinin esterase
199:Human coronavirus HKU1
1869:Microhyla letovirus 1
1677:Duck coronavirus 2714
848:Bat coronavirus HKU10
291:Genetic recombination
469:10.1128/JVI.02182-06
54:Virus classification
670:of Betacoronavirus.
457:Journal of Virology
46:electron micrograph
16:Subgenus of viruses
1327:Murine coronavirus
1300:Bovine coronavirus
1238:Canine coronavirus
1233:Feline coronavirus
1228:Alphacoronavirus 1
940:Mink coronavirus 1
785:Orthocoronavirinae
206:Murine coronavirus
39:Murine coronavirus
2014:Betacoronaviruses
1996:
1995:
1921:Taxon identifiers
1912:
1911:
1889:
1888:
1885:
1884:
1881:
1880:
1877:
1876:
1823:
1822:
1819:
1818:
1815:
1814:
1787:
1786:
1764:Coronavirus HKU15
1729:
1728:
1690:
1689:
1686:
1685:
1663:Avian coronavirus
1642:
1641:
1612:
1611:
1573:
1572:
1569:
1568:
1485:
1484:
1441:
1440:
1378:
1377:
1350:
1349:
1294:Betacoronavirus 1
1262:
1261:
1258:
1257:
1208:
1207:
1180:
1179:
1152:
1151:
1115:
1114:
1087:
1086:
1050:
1049:
1013:
1012:
985:
984:
948:
947:
920:
919:
892:
891:
864:
863:
827:
826:
733:Higher taxonomy:
530:10.3390/v11020174
221:
220:
185:Betacoronavirus 1
2021:
1989:
1988:
1976:
1975:
1963:
1962:
1961:
1948:
1947:
1946:
1916:
1915:
1904:
1862:
1861:
1853:
1852:
1844:
1843:
1829:
1828:
1800:
1799:
1742:
1741:
1714:
1713:
1703:
1702:
1697:Deltacoronavirus
1655:
1654:
1625:
1624:
1597:
1596:
1586:
1585:
1580:Gammacoronavirus
1498:
1497:
1454:
1453:
1391:
1390:
1363:
1362:
1286:
1285:
1275:
1274:
1221:
1220:
1193:
1192:
1165:
1164:
1128:
1127:
1100:
1099:
1063:
1062:
1026:
1025:
998:
997:
961:
960:
933:
932:
905:
904:
877:
876:
840:
839:
812:
811:
801:
800:
795:Alphacoronavirus
779:
778:
721:Taxonomy of the
715:
708:
701:
692:
691:
686:
682:
673:
672:
658:
640:
638:10.3390/v2081803
631:(8): 1804–1820.
616:
610:
609:
600:
571:
565:
564:
550:
532:
508:
502:
501:
488:
463:(4): 1574–1585.
448:
442:
441:
427:
409:
407:10.3390/v2081803
400:(8): 1804–1820.
385:
376:
375:
373:
371:
362:. October 2018.
352:
62:
61:
33:
19:
18:
2029:
2028:
2024:
2023:
2022:
2020:
2019:
2018:
2009:Virus subgenera
1999:
1998:
1997:
1992:
1984:
1979:
1971:
1966:
1957:
1956:
1951:
1942:
1941:
1936:
1923:
1913:
1908:
1873:
1838:
1811:
1783:
1725:
1682:
1638:
1608:
1565:
1481:
1437:
1374:
1346:
1333:Rat coronavirus
1269:Betacoronavirus
1254:
1204:
1176:
1148:
1111:
1083:
1046:
1009:
981:
944:
916:
888:
860:
823:
788:
773:
766:Cornidovirineae
754:Pisoniviricetes
727:
719:
689:
683:
676:
617:
613:
572:
568:
509:
505:
449:
445:
386:
379:
369:
367:
354:
353:
346:
342:
311:
288:
260:
239:Betacoronavirus
157:Betacoronavirus
121:Pisoniviricetes
56:
17:
12:
11:
5:
2027:
2017:
2016:
2011:
1994:
1993:
1991:
1990:
1977:
1964:
1949:
1933:
1931:
1925:
1924:
1910:
1909:
1894:
1891:
1890:
1887:
1886:
1883:
1882:
1879:
1878:
1875:
1874:
1872:
1871:
1865:
1859:
1850:
1848:Alphaletovirus
1840:
1839:
1825:
1824:
1821:
1820:
1817:
1816:
1813:
1812:
1810:
1809:
1803:
1797:
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1784:
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1781:
1774:
1767:
1760:
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1692:
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1658:
1652:
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1628:
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1614:
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1600:
1594:
1583:
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1570:
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1561:
1556:
1551:
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1541:
1536:
1531:
1526:
1525:
1524:
1514:
1501:
1495:
1487:
1486:
1483:
1482:
1480:
1479:
1472:
1465:
1457:
1451:
1443:
1442:
1439:
1438:
1436:
1435:
1428:
1421:
1420:
1419:
1414:
1402:
1394:
1388:
1380:
1379:
1376:
1375:
1373:
1372:
1366:
1360:
1352:
1351:
1348:
1347:
1345:
1344:
1337:
1336:
1335:
1323:
1316:
1309:
1308:
1307:
1302:
1289:
1283:
1272:
1264:
1263:
1260:
1259:
1256:
1255:
1253:
1252:
1251:
1250:
1245:
1240:
1235:
1224:
1218:
1210:
1209:
1206:
1205:
1203:
1202:
1196:
1190:
1182:
1181:
1178:
1177:
1175:
1174:
1168:
1162:
1154:
1153:
1150:
1149:
1147:
1146:
1139:
1131:
1125:
1117:
1116:
1113:
1112:
1110:
1109:
1103:
1097:
1089:
1088:
1085:
1084:
1082:
1081:
1074:
1066:
1060:
1052:
1051:
1048:
1047:
1045:
1044:
1037:
1029:
1023:
1015:
1014:
1011:
1010:
1008:
1007:
1001:
995:
987:
986:
983:
982:
980:
979:
972:
964:
958:
950:
949:
946:
945:
943:
942:
936:
930:
922:
921:
918:
917:
915:
914:
908:
902:
894:
893:
890:
889:
887:
886:
880:
874:
866:
865:
862:
861:
859:
858:
851:
843:
837:
829:
828:
825:
824:
822:
821:
815:
809:
798:
790:
789:
775:
774:
732:
729:
728:
718:
717:
710:
703:
695:
688:
687:
674:
611:
566:
503:
443:
377:
343:
341:
338:
337:
336:
328:
320:
310:
307:
287:
284:
259:
256:
219:
218:
217:
216:
209:
202:
195:
188:
178:
177:
173:
172:
165:
161:
160:
153:
149:
148:
141:
137:
136:
129:
125:
124:
117:
113:
112:
105:
101:
100:
93:
89:
88:
81:
74:
73:
68:
64:
63:
50:
49:
35:
34:
26:
25:
15:
9:
6:
4:
3:
2:
2026:
2015:
2012:
2010:
2007:
2006:
2004:
1987:
1982:
1978:
1974:
1969:
1965:
1960:
1954:
1950:
1945:
1939:
1935:
1934:
1932:
1930:
1926:
1922:
1917:
1907:
1903:
1898:
1892:
1870:
1867:
1866:
1863:
1860:
1858:
1854:
1851:
1849:
1845:
1841:
1837:
1836:
1830:
1826:
1808:
1805:
1804:
1801:
1798:
1796:
1795:
1790:
1780:
1779:
1775:
1773:
1772:
1768:
1766:
1765:
1761:
1759:
1758:
1754:
1752:
1751:
1747:
1746:
1743:
1740:
1738:
1737:
1732:
1722:
1719:
1718:
1715:
1712:
1710:
1709:
1704:
1701:
1699:
1698:
1693:
1679:
1678:
1674:
1672:
1671:
1667:
1665:
1664:
1660:
1659:
1656:
1653:
1651:
1650:
1645:
1635:
1634:
1630:
1629:
1626:
1623:
1621:
1620:
1615:
1605:
1602:
1601:
1598:
1595:
1593:
1592:
1591:Brangacovirus
1587:
1584:
1582:
1581:
1576:
1560:
1557:
1555:
1552:
1550:
1547:
1545:
1542:
1540:
1537:
1535:
1532:
1530:
1527:
1523:
1520:
1519:
1518:
1515:
1513:
1510:
1509:
1508:
1507:
1503:
1502:
1499:
1496:
1494:
1493:
1488:
1478:
1477:
1473:
1471:
1470:
1466:
1464:
1463:
1459:
1458:
1455:
1452:
1450:
1449:
1444:
1434:
1433:
1429:
1427:
1426:
1422:
1418:
1415:
1413:
1410:
1409:
1408:
1407:
1403:
1401:
1400:
1396:
1395:
1392:
1389:
1387:
1386:
1381:
1371:
1368:
1367:
1364:
1361:
1359:
1358:
1353:
1343:
1342:
1338:
1334:
1331:
1330:
1329:
1328:
1324:
1322:
1321:
1317:
1315:
1314:
1310:
1306:
1303:
1301:
1298:
1297:
1296:
1295:
1291:
1290:
1287:
1284:
1282:
1281:
1276:
1273:
1271:
1270:
1265:
1249:
1246:
1244:
1241:
1239:
1236:
1234:
1231:
1230:
1229:
1226:
1225:
1222:
1219:
1217:
1216:
1211:
1201:
1198:
1197:
1194:
1191:
1189:
1188:
1183:
1173:
1170:
1169:
1166:
1163:
1161:
1160:
1155:
1145:
1144:
1140:
1138:
1137:
1133:
1132:
1129:
1126:
1124:
1123:
1118:
1108:
1105:
1104:
1101:
1098:
1096:
1095:
1090:
1080:
1079:
1075:
1073:
1072:
1068:
1067:
1064:
1061:
1059:
1058:
1053:
1043:
1042:
1038:
1036:
1035:
1031:
1030:
1027:
1024:
1022:
1021:
1016:
1006:
1003:
1002:
999:
996:
994:
993:
988:
978:
977:
973:
971:
970:
966:
965:
962:
959:
957:
956:
955:Minunacovirus
951:
941:
938:
937:
934:
931:
929:
928:
923:
913:
910:
909:
906:
903:
901:
900:
895:
885:
882:
881:
878:
875:
873:
872:
871:Duvinacovirus
867:
857:
856:
852:
850:
849:
845:
844:
841:
838:
836:
835:
830:
820:
817:
816:
813:
810:
808:
807:
802:
799:
797:
796:
791:
787:
786:
780:
776:
772:
771:Coronaviridae
768:
767:
762:
761:
756:
755:
750:
749:
744:
743:
742:Orthornavirae
738:
737:
730:
726:
725:
724:Coronaviridae
716:
711:
709:
704:
702:
697:
696:
693:
681:
679:
671:
666:
662:
657:
652:
648:
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634:
630:
626:
622:
615:
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604:
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586:
582:
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570:
563:
558:
554:
549:
544:
540:
536:
531:
526:
522:
518:
514:
507:
500:
499:See figure 2.
496:
492:
487:
482:
478:
474:
470:
466:
462:
458:
454:
447:
440:
435:
431:
426:
421:
417:
413:
408:
403:
399:
395:
391:
384:
382:
365:
361:
357:
351:
349:
344:
334:
333:
329:
326:
325:
321:
318:
317:
313:
312:
306:
304:
303:African camel
300:
296:
292:
286:Recombination
283:
281:
277:
273:
269:
266:in which the
265:
255:
253:
249:
245:
244:coronaviruses
241:
240:
236:in the genus
235:
234:coronaviruses
231:
227:
226:
215:
214:
210:
208:
207:
203:
201:
200:
196:
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179:
174:
171:
170:
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145:Coronaviridae
142:
139:
138:
135:
134:
130:
127:
126:
123:
122:
118:
115:
114:
111:
110:
106:
103:
102:
99:
98:
97:Orthornavirae
94:
91:
90:
87:
86:
82:
79:
76:
75:
72:
69:
66:
65:
60:
55:
51:
47:
44:
40:
36:
32:
27:
24:
20:
1928:
1868:
1856:
1847:
1833:
1806:
1794:Herdecovirus
1792:
1776:
1769:
1762:
1755:
1748:
1736:Buldecovirus
1734:
1720:
1706:
1695:
1675:
1668:
1661:
1647:
1631:
1617:
1603:
1589:
1578:
1504:
1492:Sarbecovirus
1490:
1474:
1467:
1460:
1446:
1430:
1423:
1404:
1397:
1385:Merbecovirus
1383:
1369:
1355:
1339:
1325:
1318:
1311:
1292:
1279:
1278:
1267:
1227:
1213:
1199:
1185:
1171:
1157:
1141:
1134:
1122:Setracovirus
1120:
1106:
1094:Rhinacovirus
1092:
1076:
1069:
1055:
1039:
1032:
1020:Nyctacovirus
1018:
1004:
992:Myotacovirus
990:
974:
967:
953:
939:
925:
911:
899:Luchacovirus
897:
883:
869:
853:
846:
832:
818:
804:
793:
783:
770:
764:
758:
752:
748:Pisuviricota
746:
740:
734:
722:
668:
628:
624:
614:
580:
576:
569:
560:
520:
516:
506:
498:
460:
456:
446:
437:
397:
393:
368:. Retrieved
359:
330:
324:Merbecovirus
322:
316:Sarbecovirus
314:
289:
261:
251:
237:
224:
223:
222:
211:
204:
197:
190:
183:
168:
167:
156:
144:
132:
120:
109:Pisuviricota
108:
96:
84:
77:
67:(unranked):
22:
1959:Embecovirus
1953:Wikispecies
1929:Embecovirus
1906:Wikispecies
1857:Milecovirus
1835:Letovirinae
1708:Andecovirus
1619:Cegacovirus
1448:Nobecovirus
1357:Hibecovirus
1280:Embecovirus
1215:Tegacovirus
1187:Sunacovirus
1159:Soracovirus
1057:Pedacovirus
927:Minacovirus
834:Decacovirus
806:Colacovirus
760:Nidovirales
332:Nobecovirus
225:Embecovirus
169:Embecovirus
133:Nidovirales
23:Embecovirus
2003:Categories
1649:Igacovirus
1559:SHC014-CoV
1517:SARS-CoV-2
1512:SARS-CoV-1
523:(2): 174.
370:24 January
340:References
335:(group 2d)
327:(group 2c)
319:(group 2b)
164:Subgenus:
1944:Q57754667
736:Riboviria
647:1999-4915
539:1999-4915
477:0022-538X
416:1999-4915
299:dromedary
276:spike (S)
258:Structure
246:, have a
92:Kingdom:
85:Riboviria
1938:Wikidata
1895:Source:
1539:RacCS203
1529:BANAL-52
1522:Variants
685:31534035
665:21994708
557:30791586
495:17121802
434:21994708
364:Archived
309:See also
274:(E) and
272:envelope
268:membrane
264:envelope
230:subgenus
176:Species
140:Family:
104:Phylum:
1986:2509481
656:3185738
625:Viruses
607:9233431
598:7130985
548:6409556
517:Viruses
486:1797546
425:3185738
394:Viruses
295:genomes
152:Genus:
128:Order:
116:Class:
1973:980926
1554:RpYN06
1549:RmYN02
1544:RaTG13
663:
653:
645:
605:
595:
555:
545:
537:
493:
483:
475:
432:
422:
414:
282:(HE).
43:virion
41:(MHV)
769:>
763:>
757:>
751:>
745:>
739:>
270:(M),
228:is a
78:Realm
71:Virus
1981:NCBI
1968:BOLD
1897:ICTV
661:PMID
643:ISSN
603:PMID
553:PMID
535:ISSN
491:PMID
473:ISSN
430:PMID
412:ISSN
372:2019
1899:––
651:PMC
633:doi
593:PMC
585:doi
543:PMC
525:doi
481:PMC
465:doi
439:1).
420:PMC
402:doi
232:of
2005::
1983::
1970::
1955::
1940::
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641:.
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559:.
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519:.
515:.
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489:.
479:.
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392:.
380:^
358:.
347:^
254:.
80::
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635::
629:2
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398:2
374:.
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