Streptococcus dysgalactiae subsp. equisimilis strains (group G streptococcus [GGS]) are largely defined as commensal organisms, which are closely related to the well-defined human pathogen, the group A streptococcus (GAS). While lateral gene transfers are emerging as a common theme in these species, little is known about the mechanisms and role of these transfers and their effect on the population structure of streptococci in nature. It is now becoming evident that bacteriophages are major contributors to the genotypic diversity of GAS and, consequently, are pivotal to the GAS strain structure. Furthermore, bacteriophages are strongly associated with altering the pathogenic potential of GAS. In contrast, little is know about phages from GGS and their role in the population dynamics of GGS. In this study we report the first complete genome sequence of a GGS phage, ⌽3396. Exhibiting high homology to the GAS phage ⌽315.1, the chimeric nature of ⌽3396 is unraveled to reveal evidence of extensive ongoing genetic diversity and dissemination of streptococcal phages in nature. Furthermore, we expand on our recent findings to identify inducible ⌽3396 homologues in GAS from a region of endemicity for GAS and GGS infection. Together, these findings provide new insights into not only the population structure of GGS but also the overall population structure of the streptococcal genus and the emergence of pathogenic variants.Streptococcus dysgalactiae subsp. equisimilis (group G streptococcus [GGS]) commonly inhabits the throat, skin, and vagina of healthy humans (28). However, occasionally the organism can cause pharyngitis, impetigo, cellulitis, septicemia, glomerulonephritis, and toxic shock (14,28). This same disease spectrum generally coincides with that of a well-known human pathogen, the group A streptococcus (GAS; Streptococcus pyogenes) which cohabits the same tissue sites. Due to the extensive overlap of the disease spectrum of GGS with that caused by GAS, it is possible that disease burden specifically attributed to GGS has been historically underestimated. Of late, there are reported increases of GGS infections associated with classic human GAS-associated diseases, including invasive manifestations (10,14,15,19,23,34).Phylogenetic analyses based on rRNA sequence and other molecular clocks revealed that S. dysgalactiae spp. are closely related to S. pyogenes (16,26,35,38). The observations of overlapping clinical presentations, niche-sharing for colonization, and this evolutionary relatedness are conducive for interspecies lateral genetic transfers (LGT). Previous studies have shown that GGS possesses genes for M protein, C5a peptidase, streptokinase, streptococcal pyrogenic exotoxins, and fibronectin binding proteins (9,20,21,24,25,29,37,40). These are well defined as essential virulence determinants in GAS. The presence of mosaic structures among some of these genes within GAS and GGS strongly suggests that interspecies LGT events do indeed occur between these two streptococci and could be derived from recent and...