The flagellate Caduceia versatilis in the gut of the termite Cryptotermes cavifrons reportedly propels itself not by its own flagella but solely by the flagella of ectosymbiotic bacteria. Previous microscopic observations have revealed that the motility symbionts are flagellated rods partially embedded in the host cell surface and that, together with a fusiform type of ectosymbiotic bacteria without flagella, they cover almost the entire surface. To identify these ectosymbionts, we conducted 16S rRNA clone analyses of bacteria physically associated with the Caduceia cells. Two phylotypes were found to predominate in the clone library and were phylogenetically affiliated with the "Synergistes" phylum and the order Bacteroidales in the Bacteroidetes phylum. Probes specifically targeting 16S rRNAs of the respective phylotypes were designed, and fluorescence in situ hybridization (FISH) was performed. As a result, the "Synergistes" phylotype was identified as the motility symbiont; the Bacteroidales phylotype was the fusiform ectobiont. The "Synergistes" phylotype was a member of a cluster comprising exclusively uncultured clones from the guts of various termite species. Interestingly, four other phylotypes in this cluster, including the one sharing 95% sequence identity with the motility symbiont, were identified as nonectosymbiotic, or free-living, gut bacteria by FISH. We thus suggest that the motility ectosymbiont has evolved from a free-living gut bacterium within this termite-specific cluster. Based on these molecular and previous morphological data, we here propose a novel genus and species, "Candidatus Tammella caduceiae," for this unique motility ectosymbiont of Caducaia versatilis.Termites harbor complex gut microbiota, which comprises unicellular eukaryotes, bacteria, and archaea. The majority of the symbiotic flagellates in termite guts are characterized by both their cellulolytic activity and an abundance of prokaryotic symbionts within and/or attached to the outsides of their cell bodies. Whereas the ecological roles that these physically associated symbionts fulfill for their host protists are mostly unknown, that of ectosymbionts of the flagellate Caduceia versatilis (family Devescovinidae, order Cristamonadida, phylum Parabasalia) in the gut of the termite Cryptotermes cavifrons was clearly demonstrated by Tamm in 1982; the ectosymbiotic bacteria confer motility on the host Caduceia protists with their flagella (36). Although the host protists possess their own flagella, their rapid gliding movement is provided solely by the flagellated ectosymbiotic bacteria and never by their own flagella. This arrangement is comparable to the only other known example of motility symbiosis between prokaryotes and eukaryotic hosts, that is, between the devescovinid flagellate Mixotricha paradoxa in the gut of the termite Mastotermes darwiniensis and its ectosymbiotic spirochetes (6,19,41).Detailed microscopic observations have revealed that the motility ectosymbionts of C. versatilis are uniform rod-shaped bacteria...