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The giant colonial ciliate <jats:italic>Zoothamnium niveum<\/jats:italic>, however, is the only host of thioautotrophic symbionts that has been cultivated along with its symbiont, the vertically transmitted ectosymbiont <jats:italic>Candidatus<\/jats:italic> Thiobius zoothamnicola (short Thiobius). Because theoretical predictions posit a smaller genome in vertically transmitted endosymbionts compared to free\u2010living relatives, we investigated whether this is true also for an ectosymbiont. We used metagenomics to recover the high\u2010quality draft genome of this bacterial symbiont. For comparison we have also sequenced a closely related free\u2010living cultured but not formally described strain Milos ODIII6 (short ODIII6). We then performed comparative genomics to assess the functional capabilities at gene, metabolic pathway and trait level. 16S rRNA gene trees and average amino acid identity confirmed the close phylogenetic relationship of both bacteria. Indeed, Thiobius has about a third smaller genome than its free\u2010living relative ODIII6, with reduced metabolic capabilities and fewer functional traits. The functional capabilities of Thiobius were a subset of those of the more versatile ODIII6, which possessed additional genes for oxygen, sulphur and hydrogen utilization and for the acquisition of phosphorus illustrating features that may be adaptive for the unstable environmental conditions at hydrothermal vents. In contrast, Thiobius possesses genes potentially enabling it to utilize lactate and acetate heterotrophically, compounds that may be provided as byproducts by the host. The present study illustrates the effect of strict host\u2010dependence of a bacterial ectosymbiont on genome evolution and host adaptation.<\/jats:p>","DOI":"10.1111\/1755-0998.13889","type":"journal-article","created":{"date-parts":[[2023,11,28]],"date-time":"2023-11-28T01:17:29Z","timestamp":1701134249000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Comparative genomics of a vertically transmitted thiotrophic bacterial ectosymbiont and its close free\u2010living relative"],"prefix":"10.1111","volume":"24","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9569-409X","authenticated-orcid":false,"given":"Salvador","family":"Espada\u2010Hinojosa","sequence":"first","affiliation":[{"name":"Department of Functional and Evolutionary Ecology University of Vienna  Vienna 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