Child level

Same level

Genera

Enterovirus
Cardiovirus
Aphthovirus
Hepatovirus
Parechovirus
Erbovirus
Kobuvirus
Teschovirus
"Sapelovirus"
"Senecavirus"
"Tremovirus"
"Avihepatovirus"
Unassigned
  Genus: Cardiovirus

Species: Theilovirus

Type: Theiler's murine encephalomyelitis virus (TMEV)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Vilyuisk human encephalomyelitis virus (VHEV)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Thera virus (TRV)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Saffold virus 1 (SAFV-1)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Saffold virus 2 (SAFV-2)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Saffold virus 3 (SAFV-3)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Saffold virus 4 (SAFV-4)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Saffold virus 5 (SAFV-5)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Saffold virus 6 (SAFV-6)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Saffold virus 7 (SAFV-7)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Saffold virus 8 (SAFV-8)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR
Type: Saffold virus 9 (SAFV-9)  
5' UTR L VP4 VP2 VP3 VP1 2A 2B 2C 3A 3B 3C 3D 3' UTR

(White areas indicate genes which haven't yet been sequenced completely; grey areas are where only partial sequences exist)

Virus Strain Region Length Accession No. Reference Comments

Theiler's murine encephalomyelitis virus (TMEV)

TMEV GD VII (New York/37) 3C+3D+3' NCR 1924 M14703 Ozden et al., 1986  
CG 8105 M20562 Pevear et al., 1988  
CG 8101 X56019 Law et al., 1990  
TMEV DA (Boston/52) CG 8093 M20301 Ohara et al., 1988  
TMEV BeAn 8386 (Brazil/57) CG 8098 M16020 Pevear et al., 1987  
TMEV M2 (from BeAn 8386) L+P1 2766 AF030574 Kim et al., 1998 low-pathogenic M2 variant of strain BeAn 8386
TMEV S2 (variant of BeAn 8386) CG

8100

DQ401688

Myoung et al., 2007  
TMEV TO4 (New York/37) 5'NCR(p) 534 M80885 Pritchard et al., 1992b  
L+P1 2760 U33045 Michiels et al., 1995  
TMEV TO4 (B15) 5'NCR(p) 521 M80886 Pritchard et al., 1992b
CG 8100 EU718732 Liang et al., 2008  
TMEV FA (New York/37) 5'NCR(p) 532 M80883 Pritchard et al., 1992b  
L+P1 2766 U32924 Michiels et al., 1995  
TMEV TO Yale (1943) 5'NCR(p) 529 M80890 Pritchard et al., 1992b  
L+P1 2766 U33047 Michiels et al., 1995  
CG 8096 EU723238 Liang et al., 2008  
TMEV WW (Philadelphia/77) 5'NCR(p) 534 M80889 Pritchard et al., 1992b  
L+P1 2760 U33046 Michiels et al., 1995  
TMEV VL 5'NCR+L(p) 1100 M80887 Pritchard et al., 1992b  
TMEV ASK-1 P1 2649 U20288 Luo et al., unpublished  
TMEV ? 5'UTR(p) 58 S61105 Bandyopadhyay et al., 1993  
TMEV MHG* 5'NCR(p)+L(p)

521

M80884

Pritchard et al., 1992b  
TMEV Vie415HTR CG 8096 EU718733 Liang et al., 2008  

Vilyuisk human encephalomyelitis virus (VHEV)

VHEV V1/Siberia/55 5' NCR(p) 966 M80888 Pritchard et al., 1992b  
L+P1 2790 M94868 Pritchard et al., 1992a  
P2+P3+3'UTR 4286 EU723237 Liang et al., 2008  

Thera virus (TRV) [formerly known as Theiler-like virus (TLV) of rats]

TRV NGS910 CG

8021

AB090161

Ohsawa et al., 2003 5' end missing (~73 nt)
TRV RTV1† CG 7996 EU815052 Doel et al., unpublished 5' end missing (~100 nt)
TRV RTV1† CG 8094 EU542581 Drake et al., 2008  

Saffold virus 1 (SAFV-1)

SAFV-1 California/81 CG

8115

EF165067

Jones et al., 2007  
SAFV-1 LZ50419 5'UTR(p) 778 FJ586238 Xu et al., 2009 China: Oct-2006
VP1 896 FJ586240 Xu et al., 2009
SAFV-1 GL311
VP1

825

FJ464766 

Ren et al., 2009 China: Nov-2007
SAFV-1 GL317 VP1

825

FJ464767 Ren et al., 2009 China: Nov-2007
SAFV-1 GL328 VP1

825

FJ464768 Ren et al., 2009 China: Nov-2007
SAFV-1 GL341 VP1

825

FJ464769 Ren et al., 2009 China: Nov-2007
SAFV-1 GL352 VP1

825

FJ464770 Ren et al., 2009 China: Nov-2007
SAFV-1 GL361 VP1

825

FJ464771 Ren et al., 2009 China: Nov-2007
SAFV-1 GL362 VP1

825

FJ464772 Ren et al., 2009 China: Nov-2007
SAFV-1 GL365 VP1

825

FJ464773 Ren et al., 2009 China: Nov-2007
SAFV-1 GL368
VP1

825

FJ464774 Ren et al., 2009 China: Nov-2007
SAFV-1 GL371
VP1

825

FJ464775 Ren et al., 2009 China: Nov-2007
SAFV-1 GL376
VP1

825

FJ464776 Ren et al., 2009 China: Nov-2007
SAFV-1 GL377
VP1

825

FJ464777 Ren et al., 2009 China: Nov-2007

Saffold virus 2 (SAFV-2)

SAFV-2 Can112051-06 pp-coding
region
6882 AM922293 Abed and Boivin, 2008 Canada: 06-Mar-2006
SAFV-2 UC1 CG 7961 EU376394 Chiu et al., 2008 approx. 90 nt missing from 5' end
SAFV-2 UC3 5'UTR(p) 607 EU604742 Chiu et al., 2008  
VP1 819 EU604750 Chiu et al., 2008  
SAFV-2 UC4 5'UTR(p) 608 EU604739 Chiu et al., 2008  
VP1 819 EU604747 Chiu et al., 2008  
SAFV-2 UC6 5'UTR(p) 607 EU604744 Chiu et al., 2008  
VP1 819 EU604748 Chiu et al., 2008  
SAFV-2 UC7 5'UTR(p) 607 EU604743 Chiu et al., 2008  
VP1 819 EU604749 Chiu et al., 2008  
SAFV-2 D/VI2229/2004 P1 2532  EU681176 Drexler et al., 2008 Germany
SAFV-2 BR/118/2006 P1 2532  EU681177 Drexler et al., 2008 Brazil
SAFV-2 Pak971 VP1 816 FJ463601 Blinkova et al., 2009 Pakistan: 05-Mar-2008
2C(p) 220 FJ463607 Blinkova et al., 2009
SAFV-2 FIN08-13B VP3(p)+VP1+2A(p) 1181 FJ374267 Blomqvist et al., unpub.  

Saffold virus 3 (SAFV-3)

SAFV-3 D/VI2273/2004 P1 2538 EU681178 Drexler et al., 2008 Germany
SAFV-3 D/VI2223/2004 P1 2538 EU681179 Drexler et al., 2008 Germany
SAFV-3 UC2 5'UTR(p) 607 EU604741 Chiu et al., 2008  
VP1 813 EU604745 Chiu et al., 2008  
SAFV-3 UC5 5'UTR(p) 607 EU604740 Chiu et al., 2008  
VP1 813 EU604746 Chiu et al., 2008  
SAFV-3 Pak2678 VP1 810 FJ463605 Blinkova et al., 2009  
2C(p) 220 FJ463612 Blinkova et al., 2009
SAFV-3 Nig266 VP1(p) 723 FJ997531 Blinkova, unpub. Nigeria: March 2007
2C(p) 220 FJ997534 Blinkova, unpub.
SAFV-3 Nijmegen2007 CG 8051 FM207487 Zoll et al., 2009 Netherlands, 05-Dec-2007
SAFV-3 07-Aichi10247 VP1 813 AB542806 Ito and Yamashita, unpub. Japan: 2007
SAFV-3 07-Aichi10345 VP1 813 AB542807 Ito and Yamashita, unpub. Japan: 2007

Saffold virus 4 (SAFV-4)

SAFV-4 Pak5842 VP1 813 FJ463606 Blinkova et al., 2009  
2C(p) 220 FJ463611 Blinkova et al., 2009
SAFV-4 Pak12 VP1 813 FJ463600 Blinkova et al., 2009  
2C(p) 220 FJ463610 Blinkova et al., 2009
SAFV-4 Pak962 VP1 813 FJ463603 Blinkova et al., 2009  
2C(p) 220 FJ463608 Blinkova et al., 2009

Saffold virus 5 (SAFV-5)

SAFV-5 Pak5003 CG‡ 7627 FJ463615 Blinkova et al., 2009 ‡, 5' and 3' ends missing
SAFV-5 Pak5152 CG‡ 7639 FJ463616 Blinkova et al., 2009 ‡, 5' and 3' ends missing

Saffold virus 6 (SAFV-6)

SAFV-6 Pak6572 CG‡ 7410 FJ463617 Blinkova et al., 2009 ‡, 5' and 3' ends missing

Saffold virus 7 (SAFV-7)

SAFV-7 Afg1449 VP1 816 FJ463602 Blinkova et al., 2009 Afghanistan: 2007
2C(p) 220 FJ463613 Blinkova et al., 2009

Saffold virus 8 (SAFV-8)

SAFV-8 Pak1141 VP1 810 FJ463604 Blinkova et al., 2009 Pakistan: 13-Mar-2008
2C(p) 220 FJ463614 Blinkova et al., 2009

Saffold virus 9 (SAFV-9)

SAFV-9 Nig239 5'UTR(p)+
L+VP4+VP2(p)
653 FJ997541 Blinkova, unpub. Nigeria: March 2007
SAFV-9 Nig329? VP1(p) 789 FJ997532 Blinkova, unpub. Nigeria: March 2007
SAFV-9 Nig239 2C(p) 220 FJ997535 Blinkova, unpub. Nigeria: March 2007

Saffold virus ? (SAFV-?)

SAFV-? LZ52903 5'UTR(p) 536 FJ610244 Xu et al., 2009 China: June 2007
SAFV-? LZ53879 5'UTR(p) 538 FJ623968 Xu et al., 2009 China: 28-Sep-2007
SAFV-? LZ53010 5'UTR(p) 538 FJ586239 Xu et al., 2009 China: 28-Oct 2006
SAFV-? Pak9 2C(p) 220 FJ463609 Blinkova et al., 2009a Pakistan: 17-Apr-2008
SAFV-? Nig234 2C(p) 220 FJ997533 Blinkova, unpub. Nigeria: March 2007
SAFV-? Tun-S88 2C(p) 220 FJ997536 Blinkova, unpub. Tunisia: 2005
SAFV-? Tun-S305 2C(p) 220 FJ997537 Blinkova, unpub. Tunisia: 2005
SAFV-? Tun-S239 2C(p) 220 FJ997538 Blinkova, unpub. Tunisia: 2006
SAFV-? Tun-S90 2C(p) 220 FJ997539 Blinkova, unpub. Tunisia: 2008
SAFV-? Tun-S105 2C(p) 220 FJ997540 Blinkova, unpub. Tunisia: 2008
SAFV-? FL1-2 2C(p) 188 GQ243683 Blinkova et al., 2009b USA: 30-Nov-2007
SAFV-? FL2-3 2C(p) 188 GQ243684 Blinkova et al., 2009b USA: 01-Oct-2008
SAFV-? FL2-4 2C(p) 188 GQ243685 Blinkova et al., 2009b USA: 04-Oct-2008
SAFV-? FL2-5 2C(p) 188 GQ243686 Blinkova et al., 2009b USA: 06-Oct-2008
SAFV-? FL2-6 2C(p) 188 GQ243687 Blinkova et al., 2009b USA: 08-Oct-2008
SAFV-? FL2-7 2C(p) 188 GQ243688 Blinkova et al., 2009b USA: 11-Oct-2008
SAFV-? FL2-1 2C(p) 188 GQ243689 Blinkova et al., 2009b USA: 07-Dec-2008
SAFV-? NC-1 2C(p) 188 GQ243690 Blinkova et al., 2009b USA: 13-Nov-2007
SAFV-? CA-1 2C(p) 188 GQ243691 Blinkova et al., 2009b USA: 18-Oct-2007
             
* also known as rat encephalomyelitis virus.
†, the authors refer to this as rat theilovirus 1.
‡, 5' and 3' ends missing.
CG = complete genome; (p) = partial gene sequence.

References

Abed, Y. and Boivin, G. (2008). New Saffold cardioviruses in 3 children, Canada. Emerging Infectious Diseases 14: 834-836. 

Bandyopadhyay, P.K., Pritchard, A., Jensen, K. and Lipton, H.L. (1993). A three-nucleotide insertion in the H stem-loop of the 5' untranslated region of Theiler's virus attenuates neurovirulence. J. Virol. 67: 3691-3695.

Blinkova, O., Kapoor, A., Victoria, J., Jones, M., Wolfe, N., Naeem, A., Shaukat, S., Sharif, S., Alam, M.M., Angez, M., Zaidi, S. and Delwart, E.L. (2009a). Cardioviruses are genetically diverse and cause common enteric infections in South Asian children. J. Virol. 83: 4631-4641.

Blinkova, O., Rosario, K., Li, L., Kapoor, A., Slikas, B., Bernardin, F., Breitbart, M. and Delwart, E. (2009b). Frequent detection of highly diverse variants of cardiovirus, cosavirus, bocavirus, and circovirus in sewage samples collected in the United States. J. Clin. Microbiol. 47: 3507-3513.

Chiu, C.Y., Greninger, A.L., Kanada, K., Kwok, T., Fischer, K.F., Runckel, C., Louie, J.K., Glaser, C.A., Yagi, S., Schnurr, D.P., Haggerty, T.D., Parsonnet, J., Ganem, D. and DeRisi, J.L. (2008). Identification of cardioviruses related to Theiler's murine encephalomyelitis virus in human infections. Proc. Natl. Acad. Sci. U.S.A. 105: 14124-14129.

Dole, V.S., Banu, L.A., Perkins, C.L., Jennings, S. and Henderson, K.S. The complete nucleotide sequence of rat theilovirus 1. Unpublished. [Research Animal Diagnostic Services, Charles River, 251 Ballardvale Street, Wilmington, MA 01887, USA]

Drake, M.T., Riley, L.K. and Livingston, R.S. (2008). Differential susceptibility of SD and CD rats to a novel rat theilovirus. Comp. Med. 58: 458-464.

Drexler, J.F., de Souza Luna, L.K., Stöcker, A., Silva Almeida, P., Medrado Ribeiro, T.C., Petersen, N., Herzog, P., Pedroso, C., Huppertz, H.I., da Costa Ribeiro, H. Jr., Baumgarte, S. and Drosten, C. (2008). Circulation of 3 lineages of a novel saffold cardiovirus in humans. Emerg. Infect. Dis. 14: 1398-1405.

Jones, M.S., Lukashov, V.V., Ganac, R.D., Schnurr, D.P. (2007). Discovery of a novel human picornavirus from a pediatric patient presenting with fever of unknown origin. J. Clin. Microbiol. 45: 2144-2150.

Kim, B.S., Yauch, R.L., Bahk, Y.Y., Kang, J.A., Dal Canto, M.C. and Hall, C.K. (1998). A spontaneous low-pathogenic variant of Theiler's virus contains an amino acid substitution within the predominant VP1(233-250) T-cell epitope. J. Virol. 72: 1020-1027. Erratum in: J. Virol 1998 72: 6965.

Law, K.M. and Brown, T.D.K. (1990). The complete nucleotide sequence of the GDVII strain of Theiler's murine encephalomyelitis virus (TMEV). Nucl. Acids Res. 18: 6707-6708.

Liang, Z., Manoj Kumar, A.S., Jones, M.S., Knowles, N.J. and Lipton, H.L. (2008). Phylogenetic analysis of the species Theilovirus: emerging murine and human pathogens. Journal of Virology 82: 11545–11554.

Luo, M., Pritchard, A.E. and Lipton, H. The structure of a highly virulent Theiler's murine encephalomyelitis virus (GDVII strain) and implications for determinants of viral persistence. Unpublished. (Pritchard A. E., University Colorado Health Sciences Center, Biochemistry, Biophysics and Genetics, 4200 E. Ninth, Denver, CO 80262, USA)

Michiels, T., Jarousse, N. and Brahic, M. (1995). Analysis of the leader and capsid coding regions of persistent and neurovirulent strains of Theiler's virus. Virology 214: 550-558.

Myoung, J., Hou, W., Kang, B., Lyman, M.A., Kang, J.-A. and Kim, B.S. (2007). The immunodominant CD8+ T cell epitope region of Theiler's virus in resistant C57BL/6 mice is critical for anti-viral immune responses, viral persistence, and binding to the host cells. Virology 360: 159-171.

Ohara, Y., Stein, S., Fu, J., Stillman, L., Klaman, L. and Roos, R.P. (1988). Molecular cloning and sequence determination of DA strain of Theiler's murine encephalomyelitis viruses. Virology 164: 245-255.

Ohsawa, K., Watanabe, Y., Miyata, H., Sato, H. (2003). Genetic analysis of a Theiler-like virus isolated from rats. Comp Med. 53: 191-196.

Ozden, S., Tangy, F., Chamorro, M. and Brahic, M. (1986). Theiler's virus genome is closely related to that of encephalomyocarditis virus, the prototype cardiovirus. J. Virol. 60: 1163-1165.

Pevear, D.C., Calenoff, M., Rozhon, E. and Lipton, H.L. (1987). Analysis of the complete nucleotide sequence of the picornavirus Theiler's murine encephalomyelitis virus indicates that it is closely related to cardioviruses. J. Virol. 61: 1507-1516.

Pevear, D.C., Borkowski, J., Calenoff, M., Oh, C.K., Ostrowski, B. and Lipton, H.L. (1988). Insights into Theiler's virus neurovirulence based on a genomic comparison of the neurovirulent GDVII and less virulent BeAn strains. Virology 165: 1-12.

Pritchard, A.E., Strom, T., and Lipton, H.L. (1992a). Nucleotide sequence identifies vilyuisk virus as a divergent Theiler's virus. Virology 191: 469-472.

Pritchard, A.E., Calenoff, M.A., Simpson, S., Jensen, K., and Lipton, H.L. (1992b). A single base deletion in the 5' noncoding region of Theiler's virus attenuates neurovirulence. J. Virol. 66: 1951-1958.

Ren, L., Gonzalez, R., Xiao, Y., Xu, X., Chen, L., Vernet, G., Paranhos-Baccala, G., Jin, Q. and Wang, J. (2009). Saffold cardiovirus in children with acute gastroenteritis, Beijing, China. Emerging Infect. Dis. 15: 1509-1511.

Xu, Z.Q., Cheng, W.X., Qi, H.M., Cui, S.X., Jin, Y. and Duan, Z.J. (2009). New Saffold cardiovirus in children, China. Emerging Infect. Dis. 15: 993-994.

Zoll, J., Erkens-Hulshof, S., Lanke, K., Verduyn-Lunel, F., Melchers, W.J.G., Schoondermark-van de Ven, E., Roivainen, M., Galama, J.M.D.,  and van Kuppeveld, F.J.M. (2009). Saffold virus, a human Theiler's-like cardiovirus, is ubiquitous and causes infection early in life. PLoS Pathog. 5(5):e1000416-e1000416.

 

 
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