Tomato spotted wilt virus (TSWV) causes severe economic losses in many crops worldwide and often overcomes resistant cultivars used for disease control. Comparison of nucleotide and amino acid sequences suggested that tomato resistance conferred by the gene Sw-5 can be overcome by the amino acid substitution C to Y at position 118 (C118Y) or T120N in the TSWV movement protein, NSm. Phylogenetic analysis revealed that substitution C118Y has occurred independently three times in the studied isolates by convergent evolution, whereas the substitution T120N was a unique event. Analysis of rates of non-synonymous and synonymous changes at individual codons showed that substitution C118Y was positively selected.Tomato spotted wilt virus (TSWV) is one of the most economically important plant viruses and causes damage in many agronomic crops worldwide (Adkins, 2000). TSWV is the type member of the genus Tospovirus, which contains the only plant-infecting members of the family Bunyaviridae. The genome consists of three negative-sense or ambisense RNA segments: segment L encodes a putative RNAdependent RNA polymerase (de Haan et al., 1991); segment M encodes the cell-to-cell movement protein, NSm (Li et al., 2009), and the precursor of the surface glycoproteins, G N / G C , involved in TSWV transmission by thrips (Sin et al., 2005;Naidu et al., 2008); and segment S encodes a silencing suppressor, NSs (Takeda et al., 2002), and the nucleocapsid, N (de Haan et al., 1990).Controlling TSWV in crop plants has proven difficult because of the wide range of plant hosts (.1000 species) and effective spread of TSWV by thrip vectors (Hanssen et al., 2010). Intense efforts have been made around the world to obtain genetically resistant cultivars against TSWV. Only the dominant genes Sw-5 and Tsw introgressed in tomato (Lycopersicon esculentum, 5Solanum lycopersicum) and pepper (Capsicum annuum) cultivars, respectively, have been found to confer resistance to a wide spectrum of TSWV isolates; they have been deployed in commercial cultivars worldwide. Sw-5 is also effective against two other tospoviruses, Tomato chlorotic spot virus (TCSV) and Groundnut ringspot virus (GRSV) (Soler et al., 2003). However, Sw-5 resistance-breaking (SRB) isolates have been detected in Hawaii, Australia, South Africa, Spain and Italy (Cho et al., 1996;Thompson & van Zijl, 1996;Latham & Jones, 1998;Aramburu & Martí, 2003;Ciuffo et al., 2005) and Tsw resistance-breaking (TRB) isolates have been detected in Brazil, USA, Italy, Spain and Australia (Boiteux et al., 1993;Hobbs et al., 1994;Roggero et al., 2002; Margaria et al., 2004;Sharman & Persley, 2006).Understanding the molecular mechanisms linked to resistance breakdown and the evolutionary processes involved in the emergence of resistance-breaking isolates is a major challenge. This is relevant to the development of more durable and efficient resistance, which would certainly have a considerable economic impact on agriculture. Since a reverse-genetics system based on infectious cDNA clones is not availab...