We describe here the fourth listerial membrane-damaging virulence factor, a
sphingomyelinase C (SMase) that is produced specifically by the ruminant p
athogen Listeria ivanovii. Its coding gene, smcL, is a monocistron expresse
d independently of PrfA. The smcL product, SmcL, is highly similar to the s
taphylococcal B-toxin and is responsible for the differential hemolytic pro
perties of L. ivanovii (bizonal hemolysis and CAMP-like reaction with R. eq
ui). The role of SmcL in virulence was assessed by gene disruption and comp
lementation. Our data show that SmcL mediates disruption of the membrane of
primary phagosomes, thereby promoting bacterial intracellular proliferatio
n. They also suggest that SmcL may play a role in host tropism. smcL is loc
ated in LIPI-2, a novel 18-kb pathogenicity island which also contains a cl
uster of internalin genes. LIPI-2 is unstable, L. ivanovii-specific and req
uired for full virulence in mice and lambs.