The chonotrichs are sessile ciliated protozoa that are ectosymbiotic on the body parts of a variety of crustaceans. They have long been considered a separate group because their sessile habit has resulted in the evolution of a very divergent body form and reproductive strategy compared to free-living ciliates. In the mid-20th Century, the free-living dysteriid cyrtophorian ciliates were proposed as a potential sister clade because the chonotrich bud or daughter cell showed similarities during division morphogenesis (i.e. ontogeny) to these free-living dysteriids. A single small subunit (SSU) rRNA gene sequence is available for the chonotrich Isochona sp. However, its authenticity has recently been questioned, and the placement of this sequence within the dysteriid clade has added to this controversy. In this report, the SSUrRNA gene sequence of the chonotrich Chilodochona carcini, ectosymbiotic on the green crab Carcinus maenas, is provided. Topology testing of the SSUrRNA gene phylogeny, constructed by Bayesian Inference, robustly supports the sister-group relationship of Isochona sp. and Chilodochona carcini, the monophyly of these two chonotrichs, and the divergence of the chonotrich clade within the dysteriid clade.
INTRODUCTIONThe chonotrichs, which are sessile symbionts, typically on the mouthparts of crustaceans, have long been recognized as a group of ciliates, although their position within the phylum was not stable until the late 20th Century (Corliss, 1979). Like the sessile suctorians and peritrichs, chonotrichs were separated out from other more motile groups at a high taxonomic rank, primarily because of their sessile nature and the unusual morphology associated with their sessility. Guilcher (1951) carefully studied the morphology of the dispersive/divisional stage of the chonotrichs, the bud or swarmer, and recognized homologies with free-living cyrtophorian ciliates, a conclusion confirmed by, among others, Dobroza nska-Kaczanowska (1963). Grain & Batisse (1974) examined the cortical microtubular components of Chilodochona and found strong similiarities to the cortical components of free-living cyrtophorians, and Fahrni (1982) provided similar data for the cortex of Spirochona gemmipara. These observations strengthened further the hypothesis that the chonotrichs arose from a free-swimming cyrtophorian, perhaps a dysteriid-like ancestor. Ultrastructural examination of the attachment apparatus of chonotrichs also showed significant similarities to the attachment apparatus of dysteriid ciliates (Fahrni 1984;Faur e-Fremiet et al., 1968; Faur e-Fremiet et al., 1956). Snoeyenbos-West et al. (2004) sequenced the small subunit (SSU) rRNA genes of several phyllopharyngean ciliates, including a species of the chonotrich genus Isochona. The Isochona gene sequence robustly clustered within the cyrtophorian clade, sister to the genes of Dysteria species. While Snoeyenbos-West et al. (2004) provided photographic evidence for their identification of the free-living cyrtophorians, Chlamydodon and Dysteria, an...