{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T10:25:02Z","timestamp":1786962302525,"version":"build-2736575974"},"posted":{"date-parts":[[2026,8,13]]},"group-title":"Microbiology","reference-count":157,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T00:00:00Z","timestamp":1786579200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"award":["#9326"],"award-info":[{"award-number":["#9326"]}],"id":[{"id":"https:\/\/ror.org\/006wxqw41","id-type":"ROR","asserted-by":"publisher"}]},{"name":"National Science Foundation","award":["DEB-2144304"],"award-info":[{"award-number":["DEB-2144304"]}]},{"name":"National Science Foundation","award":["OPP-2132641"],"award-info":[{"award-number":["OPP-2132641"]}]},{"id":[{"id":"https:\/\/ror.org\/04sqkw483","id-type":"ROR","asserted-by":"publisher"}]},{"name":"National Institute of Health National Institute of General Medical Sciences","award":["1T32GM142623"],"award-info":[{"award-number":["1T32GM142623"]}]},{"name":"Joint Genome Institute","award":["505025"],"award-info":[{"award-number":["505025"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2026,8,13]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:p>\n                  Microbial symbioses drive the evolutionary and functional diversification of eukaryotic clades, from single-celled protists to large invertebrates. However, our knowledge of host-associated assemblages (the \u201cholobiont\u201d) is limited in microscopic animal phyla with a body size &lt;1mm, due to practical challenges such as low biomass and difficult taxonomy of host species. Marine nematodes represent an ideal case study for rapidly advancing our knowledge of bacterial-animal symbioses, representing a globally abundant invertebrate group with strong links to terrestrial and model organism species within the same phylum. Here, we sequenced the holobionts of 220 marine nematodes and generated 815 metagenome-assembled genomes (MAGs) of host-associated bacteria\/archaea. Our data indicates that 20-34% of marine nematodes harbor an obligate intracellular symbiont, often with multiple endosymbionts co-occurring within the same host. Three bacterial phyla (Pseudomonadota Bacteroidota, and Verrucomicrobiota) account for three-quarters of all nematode-associated MAGs, and the majority of these holobiont MAGs represent deeply divergent lineages in the prokaryotic tree of life. The Flavobacteriaceae (a core microbiome taxon in\n                  <jats:italic>C. elegans<\/jats:italic>\n                  and other terrestrial nematodes), were consistently recovered across phylogenetically diverse marine nematode lineages, suggesting evolutionary conservation of holobiont taxa across marine and terrestrial environments. We also report a novel chemoautotroph family (\n                  <jats:italic>Ca.<\/jats:italic>\n                  Thionematobacter) recovered from nematode hosts in both deep-sea and shallow-water habitats, and report the first confirmed instance of\n                  <jats:italic>Cardinium<\/jats:italic>\n                  endosymbionts from marine invertebrates. Finally, \u223c65% of nematode-associated MAGs are able to degrade chitin, via hexosaminidase, implying that benthic invertebrate holobionts make significant contributions to global carbon cycling. These results underline the importance of evaluating symbiosis in microscopic marine invertebrates, and accelerating our understanding of animal evolution and ecosystem dynamics in vast benthic habitats.\n                <\/jats:p>","DOI":"10.64898\/2026.08.12.744518","type":"posted-content","created":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T04:50:39Z","timestamp":1786683039000},"source":"Crossref","is-referenced-by-count":0,"title":["Marine nematodes exhibit widespread symbiosis, novel chemoautotrophy, and evolutionary conservation of holobiont taxa"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9086-3050","authenticated-orcid":false,"given":"Alejandro","family":"De Santiago","sequence":"first","affiliation":[{"name":"Institute of Bioinformatics, University of Georgia, Athens GA 30605"},{"name":"Department of Marine Sciences, University of Georgia, Athens, GA 30605"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-9058-100X","authenticated-orcid":false,"given":"Min Khant","family":"Han","sequence":"additional","affiliation":[{"name":"Department of Marine Sciences, University of Georgia, Athens, GA 30605"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0004-8358-5725","authenticated-orcid":false,"given":"Sarah B.","family":"Hargadon","sequence":"additional","affiliation":[{"name":"Department of Marine Sciences, University of Georgia, Athens, GA 30605"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9417-7867","authenticated-orcid":false,"given":"Mirayana","family":"Marcellino Barros","sequence":"additional","affiliation":[{"name":"Department of Marine Sciences, University of Georgia, Athens, GA 30605"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Simone","family":"Brito De Jesus","sequence":"additional","affiliation":[{"name":"Department of Marine Sciences, University of Georgia, Athens, GA 30605"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6424-2848","authenticated-orcid":false,"given":"Tiago Jos\u00e9","family":"Pereira","sequence":"additional","affiliation":[{"name":"Institute of Bioinformatics, University of Georgia, Athens GA 30605"},{"name":"Department of Marine Sciences, University of Georgia, Athens, GA 30605"},{"name":"Department of Ecology, Evolution, and Organismal Biology, Kennesaw State University, Kennesaw, GA 30144"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4356-3837","authenticated-orcid":false,"given":"Holly M.","family":"Bik","sequence":"additional","affiliation":[{"name":"Institute of Bioinformatics, University of Georgia, Athens GA 30605"},{"name":"Department of Marine Sciences, University of Georgia, Athens, GA 30605"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"issue":"12","key":"2026081702350651000_2026.08.12.744518v1.1","doi-asserted-by":"crossref","first-page":"3479","DOI":"10.3390\/ani11123479","article-title":"Twenty years after DE Ley and Blaxter-how far did we progress in understanding the phylogeny of the phylum Nematoda?","volume":"11","year":"2021","journal-title":"Animals: An Open Access Journal from MDPI"},{"key":"2026081702350651000_2026.08.12.744518v1.2","doi-asserted-by":"publisher","DOI":"10.3389\/fevo.2021.769565"},{"key":"2026081702350651000_2026.08.12.744518v1.3","doi-asserted-by":"publisher","DOI":"10.1016\/0278-4343(90)90034-J"},{"key":"2026081702350651000_2026.08.12.744518v1.4","doi-asserted-by":"publisher","DOI":"10.1128\/AEM.56.6.1919-1925.1990"},{"key":"2026081702350651000_2026.08.12.744518v1.5","unstructured":"Andrews, S. , & Others. 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