Spiroplasmas: evolutionary relationships and biodiversity.

Spiroplasmas are wall-less descendants of Gram-positive bacteria that maintain some of the smallest genomes known for self-replicating organisms. These helical, motile prokaryotes exploit numerous habitats, but are most often found in association with insects. Co-evolution with their insect hosts may account for the highly speciose nature of the genus Spiroplasma, with many spiroplasmas existing in obligate insect/plant transmission cycles. In addition to insect and plant hosts, spiroplasmas are found in association with ticks and crustaceans. Although most spiroplasma associations appear to be commensal, some cases of pathogenicity or mutualism have been described. Most notably, spiroplasmas have been identified as the causative agents of agricultural and aquacultural diseases and the sex ratio disorder in insects. Some spiroplasmas exhibit strict host and/or geographical ranges, but others are relative generalists. Species of the genus Spiroplasma have been traditionally classified into 34 groups based on cross-reactivity of surface antigens. Three of the serogroups contain closely related strain complexes that are further divided into subgroups. Phylogenetic reconstructions based on 16S rDNA sequence strongly support the closely related serogroups. To date, less than 40 Spiroplasma species have been fully characterized and given binomial names. Complete characterization of a new species involves numerous phenotypic and genotypic tests as outlined in the minimal standards document; this document is currently under revision to include phylogenetic data and a reevaluated set of required phenotypic and genotypic tests. The area of spiroplasma research is poised for major advances with new criteria for naming species in preparation, a dramatic increase in available molecular characters, the promise of full genome sequences, and advances in genetic tools for manipulation of these organisms.