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This group is unified by the ability to form sheaths and oxidize metals, although\n                      <jats:italic toggle=\"yes\">L. ochracea<\/jats:italic>\n                      exhibits obvious ecological, morphological, and functional differences from the rest of\n                      <jats:italic toggle=\"yes\">Sphaerotilus-Leptothrix<\/jats:italic>\n                      . Recently, there have been calls to combine the group into one genus,\n                      <jats:italic toggle=\"yes\">Sphaerotilus<\/jats:italic>\n                      ; however, these studies lacked adequate genomic representation of\n                      <jats:italic toggle=\"yes\">L. ochracea<\/jats:italic>\n                      . Here, we present a comprehensive comparative genomic analysis of the\n                      <jats:italic toggle=\"yes\">Sphaerotilus-Leptothrix<\/jats:italic>\n                      group, including expanded representation of\n                      <jats:italic toggle=\"yes\">L. ochracea<\/jats:italic>\n                      , a closely related novel species,\n                      <jats:italic toggle=\"yes\">Leptothrix toolikensis<\/jats:italic>\n                      , and two new isolates (\n                      <jats:italic toggle=\"yes\">Leptothrix mechoopdaensis<\/jats:italic>\n                      ). Analysis of 38 genomes resolves three phylogenetic and functional groups: the\n                      <jats:italic toggle=\"yes\">ochracea<\/jats:italic>\n                      -type\n                      <jats:italic toggle=\"yes\">Leptothrix<\/jats:italic>\n                      (Group 1), the\n                      <jats:italic toggle=\"yes\">mobilis<\/jats:italic>\n                      -type\n                      <jats:italic toggle=\"yes\">Leptothrix<\/jats:italic>\n                      (Group 2), and\n                      <jats:italic toggle=\"yes\">Sphaerotilus<\/jats:italic>\n                      (Group 3). Group 1 genomes form a separate genus based on average nucleotide identity and alignment fraction. The genomes clearly diverge from the rest of\n                      <jats:italic toggle=\"yes\">Sphaerotilus-Leptothrix<\/jats:italic>\n                      in phylogeny, size, and metabolic potential. Group 1 genomes are much smaller (2.59\u20133.04 Mb) than those of Groups 2 (4.55\u20136.06 Mb) and 3 (3.94\u20135.07 Mb), while encoding more metal oxidases and fewer carbohydrate-active enzymes. Group 2 clusters with Group 3 phylogenetically and is similar in organic carbon metabolisms but maintains more metal oxidation genes. Group 2 members lack homogeneity in phenotype and genotype, suggesting that additional isolates and genomes are needed for confident classification. However, Group 1 genomes (\n                      <jats:italic toggle=\"yes\">L. ochracea<\/jats:italic>\n                      and\n                      <jats:italic toggle=\"yes\">L. toolikensis<\/jats:italic>\n                      ) show clear divergence, precluding their inclusion in\n                      <jats:italic toggle=\"yes\">Sphaerotilus<\/jats:italic>\n                      and supporting the retention of the genus\n                      <jats:italic toggle=\"yes\">Leptothrix<\/jats:italic>\n                      .\n                    <\/jats:p>\n                    <jats:sec>\n                      <jats:title>IMPORTANCE<\/jats:title>\n                      <jats:p>\n                        Researchers have long noted differences in metal oxidation, morphology, and ecology among\n                        <jats:italic toggle=\"yes\">Sphaerotilus-Leptothrix<\/jats:italic>\n                        , but longstanding confusion over phylogeny and genus boundaries led to inconsistent taxonomic classification between the two genera. This confusion stems from previous work that used isolates that are unavailable or lost distinguishing traits in culture, and from limited genomic data. Furthermore, the\n                        <jats:italic toggle=\"yes\">Leptothrix<\/jats:italic>\n                        type strain\n                        <jats:italic toggle=\"yes\">L. ochracea<\/jats:italic>\n                        has never been isolated. This study provides molecular evidence that substantiates calls to reassign some\n                        <jats:italic toggle=\"yes\">Leptothrix<\/jats:italic>\n                        members to the genus\n                        <jats:italic toggle=\"yes\">Sphaerotilus<\/jats:italic>\n                        but adds to an emerging body of evidence that Group 1\n                        <jats:italic toggle=\"yes\">L. ochracea<\/jats:italic>\n                        and now\n                        <jats:italic toggle=\"yes\">L. toolikensis<\/jats:italic>\n                        represent a functionally distinct lineage. While genomic similarity metrics left taxonomic divisions unclear, integrating metabolic potential with phylogeny resolved genus boundaries based on clear functional groupings. This polyphasic approach for delineating genera clarifies longstanding taxonomic confusion and refines our understanding of functional diversity both across and within\n                        <jats:italic toggle=\"yes\">Sphaerotilus-Leptothrix<\/jats:italic>\n                        lineages.\n                      <\/jats:p>\n                    <\/jats:sec>\n                  <\/jats:sec>","DOI":"10.1128\/msystems.01768-25","type":"journal-article","created":{"date-parts":[[2026,6,12]],"date-time":"2026-06-12T13:01:09Z","timestamp":1781269269000},"update-policy":"https:\/\/doi.org\/10.1128\/asmj-crossmark-policy-page","source":"Crossref","is-referenced-by-count":1,"title":["Distinguishing\n                    <i>Leptothrix<\/i>\n                    and\n                    <i>Sphaerotilus<\/i>\n                    genera by an integrated genomic-phenotypic analysis supported by new\n                    <i>Leptothrix<\/i>\n                    genomes"],"prefix":"10.1128","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7792-3880","authenticated-orcid":true,"given":"Gracee K.","family":"Tothero","sequence":"first","affiliation":[{"name":"Delaware Biotechnology Institute","place":["Newark, USA"]},{"id":[{"id":"https:\/\/ror.org\/01sbq1a82","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Earth Sciences, University of Delaware","place":["Newark, USA"]},{"id":[{"id":"https:\/\/ror.org\/01sbq1a82","id-type":"ROR","asserted-by":"publisher"}],"name":"Microbiology Graduate Program, University of Delaware","place":["Newark, USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0302-3588","authenticated-orcid":true,"given":"Jessica L.","family":"Keffer","sequence":"additional","affiliation":[{"name":"Delaware Biotechnology Institute","place":["Newark, USA"]},{"id":[{"id":"https:\/\/ror.org\/01sbq1a82","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Earth Sciences, University of Delaware","place":["Newark, USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1737-356X","authenticated-orcid":false,"given":"David","family":"Emerson","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/03v2r6x37","id-type":"ROR","asserted-by":"publisher"}],"name":"Bigelow Laboratory for Ocean Sciences","place":["East Boothbay, USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-2944-5575","authenticated-orcid":false,"given":"Emily J.","family":"Fleming","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/027bzz146","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Biological Sciences, California State University","place":["Chico, USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1810-4994","authenticated-orcid":true,"given":"Clara S.","family":"Chan","sequence":"additional","affiliation":[{"name":"Delaware Biotechnology Institute","place":["Newark, USA"]},{"id":[{"id":"https:\/\/ror.org\/01sbq1a82","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Earth Sciences, University of Delaware","place":["Newark, USA"]},{"id":[{"id":"https:\/\/ror.org\/01sbq1a82","id-type":"ROR","asserted-by":"publisher"}],"name":"Microbiology Graduate Program, University of Delaware","place":["Newark, USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"235","reference":[{"key":"e_1_3_4_2_2","doi-asserted-by":"publisher","DOI":"10.5962\/bhl.title.124909"},{"key":"e_1_3_4_3_2","first-page":"1","article-title":"Eisenbakterien als anorgoxydanten","volume":"57","author":"Winogradsky S","year":"1922","unstructured":"Winogradsky S. 1922. 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