Abstract

ABSTRACTSix subspecies are currently recognized in Salmonella enterica. Subspecies I (subspecies enterica) is responsible for nearly all infections in humans and warm-blooded animals, while five other subspecies are isolated principally from cold-blooded animals. We sequenced 21 phylogenetically diverse strains, including two representatives from each of the previously unsequenced five subspecies and 11 diverse new strains from S. enterica subspecies enterica, to put this species into an evolutionary perspective. The phylogeny of the subspecies was partly obscured by abundant recombination events between lineages and a relatively short period of time within which subspeciation took place. Nevertheless, a variety of different tree-building methods gave congruent evolutionary tree topologies for subspeciation. A total of 285 gene families were identified that were recruited into subspecies enterica, and most of these are of unknown function. At least 2,807 gene families were identified in one or more of the other subspecies that are not found in subspecies I or Salmonella bongori. Among these gene families were 13 new candidate effectors and 7 new candidate fimbrial clusters. A third complete type III secretion system not present in subspecies enterica (I) isolates was found in both strains of subspecies salamae (II). Some gene families had complex taxonomies, such as the type VI secretion systems, which were recruited from four different lineages in five of six subspecies. Analysis of nonsynonymous-to-synonymous substitution rates indicated that the more-recently acquired regions in S. enterica are undergoing faster fixation rates than the rest of the genome. Recently acquired AT-rich regions, which often encode virulence functions, are under ongoing selection to maintain their high AT content.

Highlights

  • Six subspecies are currently recognized in Salmonella enterica

  • Salmonella is considered to consist of two species, Salmonella bongori and Salmonella enterica, and S. enterica is further classified into six subspecies, arizonae (IIIa), diarizonae (IIIb), houtenae (IV), salamae (II), indica (VI), and enterica (I) [7]

  • Gene families gained at the subspecies enterica (I) node may provide clues to the strategies and virulence factors that contributed to the formation of a lineage which has evolved to infect principally warm-blooded hosts

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Summary

Introduction

Six subspecies are currently recognized in Salmonella enterica. Subspecies I (subspecies enterica) is responsible for most infections in humans and warm-blooded animals, while five other subspecies are isolated principally from coldblooded animals. IMPORTANCE We have sequenced 21 new genomes which encompass the phylogenetic diversity of Salmonella, including strains of the previously unsequenced subspecies arizonae, diarizonae, houtenae, salamae, and indica as well as new diverse strains of subspecies enterica. We have sequenced the genomes of eleven additional members of subspecies enterica (I), selected based on their diversity, and those of two different serovars from each of the other five known subspecies. Gene families gained at the subspecies enterica (I) node may provide clues to the strategies and virulence factors that contributed to the formation of a lineage which has evolved to infect principally warm-blooded hosts. We identified gene families undergoing accelerated evolution based on pairwise synonymous-tononsynonymous single nucleotide polymorphism (SNP) ratios This group of gene families is interesting because some of this selection may be driven by newly acquired life strategies or by interactions of the bacterium with the host

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