The evolution of mitogenomic architecture in Bilateria | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article The evolution of mitogenomic architecture in Bilateria Ivan Jakovlic, Dong Zhang, Yiwen Ma, Tong Ye, Rui Han, Xiang Liu, and 7 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6693840/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 12 Mar, 2026 Read the published version in Nature Communications → Version 1 posted You are reading this latest preprint version Abstract We used a dataset comprising over ten thousand bilaterian mitogenomes to address multiple open questions regarding the evolution of mitogenomic architecture. Double-stranded architecture is the most likely ancestral state for all major radiations (phylum and above), aside from Kinorhyncha, Syndermata and Brachiopoda. We found over twenty transitions from double-stranded to single-stranded (all genes on a single strand) mitogenomic architectures, as well as the first evidence of single-stranded to double-stranded transitions: one in Annelida and two or more in Mollusca. Gene order rearrangement rates (GORR) were strongly positively correlated with sequence evolution rates (r=0.69, R2≈35%). Both variables were increased in single-stranded mitogenomes, parasitic species, and those with low locomotory capacity. The latter two categories also exhibited increased proportions of single-stranded mitogenomes. Contrary to this, mitogenome size exhibited a negative correlation with both GORR and sequence evolutionary rates. Purifying selection plays a major role in the mitogenomic architectural evolution, but other variables need to be invoked to fully explain the observed patterns. Biological sciences/Evolution/Molecular evolution Earth and environmental sciences/Ecology/Evolutionary ecology mtDNA mitochondrion gene order rearrangement parasitism metabolic rate molecular evolution purifying selection locomotion strand animals Full Text Additional Declarations There is NO Competing Interest. Supplementary Files Supplementaryfile.pdf Supplementary file Supplementaryfile2.xlsx Dataset 1 rs2.pdf Reporting Summary Cite Share Download PDF Status: Published Journal Publication published 12 Mar, 2026 Read the published version in Nature Communications → Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6693840","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":471998001,"identity":"00762502-29be-4221-84bb-e106442119a4","order_by":0,"name":"Ivan Jakovlic","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA7klEQVRIiWNgGAWjYBACNhAhwZDAwMDO2MDwgQHEIloLM2MD4wxitEABSAsQ8RCjhU8i+dkDi5o0OYPDzI2fbdvs8vjZGxg/fMzB4zCJNHMDiWM5xgaHGZulc9uSiyV7DjBLztyGRwvPATMJCbaKxJnNjG3MuW3MiRtuJLAx8+LVcvybhMQ/qBbLtnoitLD3mElItuUk9jMDtTC2HSZKS5mEZF+aMT8zY7Nkz7njiTN7Djbj9Yt8M/s2aYlvyXJs7O0PP/woq07sZ28++OEjHi0gwCwBYzGCYxaYDAgBxg9w5h+CikfBKBgFo2AEAgA9oEmCnbvPUAAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0002-2461-3712","institution":"Lanzhou University","correspondingAuthor":true,"prefix":"","firstName":"Ivan","middleName":"","lastName":"Jakovlic","suffix":""},{"id":471998002,"identity":"5a802426-9eaf-4af0-9c4e-b81f4a1534d6","order_by":1,"name":"Dong Zhang","email":"","orcid":"https://orcid.org/0000-0002-0902-6704","institution":"Lanzhou University","correspondingAuthor":false,"prefix":"","firstName":"Dong","middleName":"","lastName":"Zhang","suffix":""},{"id":471998003,"identity":"20dedd9d-a613-4e2e-86d0-a5bf8bfb1329","order_by":2,"name":"Yiwen Ma","email":"","orcid":"","institution":"Lanzhou University","correspondingAuthor":false,"prefix":"","firstName":"Yiwen","middleName":"","lastName":"Ma","suffix":""},{"id":471998004,"identity":"f5bbb8c4-bd4e-483b-9f39-f9a44c184215","order_by":3,"name":"Tong Ye","email":"","orcid":"","institution":"Lanzhou University","correspondingAuthor":false,"prefix":"","firstName":"Tong","middleName":"","lastName":"Ye","suffix":""},{"id":471998005,"identity":"4464ab2b-a28e-4ae6-9484-dcf9dbad5f1a","order_by":4,"name":"Rui Han","email":"","orcid":"","institution":"State Key Laboratory of Herbage Improvement and Grassland Agro-ecosystems, and College of Ecology, Lanzhou University","correspondingAuthor":false,"prefix":"","firstName":"Rui","middleName":"","lastName":"Han","suffix":""},{"id":471998006,"identity":"609a0c19-de18-4a5b-b1ad-f149652eb1a7","order_by":5,"name":"Xiang Liu","email":"","orcid":"https://orcid.org/0000-0002-9422-9492","institution":"Lanzhou University","correspondingAuthor":false,"prefix":"","firstName":"Xiang","middleName":"","lastName":"Liu","suffix":""},{"id":471998007,"identity":"9cc92801-bea9-4ef9-874b-ad59af744044","order_by":6,"name":"Hong Zou","email":"","orcid":"","institution":"Chinese Academy of Sciences","correspondingAuthor":false,"prefix":"","firstName":"Hong","middleName":"","lastName":"Zou","suffix":""},{"id":471998008,"identity":"9655ff59-19a4-4bca-862c-b00a856e8fdd","order_by":7,"name":"Jin Zhang","email":"","orcid":"","institution":"Bio-Transduction Lab","correspondingAuthor":false,"prefix":"","firstName":"Jin","middleName":"","lastName":"Zhang","suffix":""},{"id":471998009,"identity":"13459287-b56f-481e-9696-654f59839020","order_by":8,"name":"Rui Song","email":"","orcid":"","institution":"Hunan Fisheries Research Institute","correspondingAuthor":false,"prefix":"","firstName":"Rui","middleName":"","lastName":"Song","suffix":""},{"id":471998010,"identity":"1e267af9-598f-462d-9cae-ea90157105a4","order_by":9,"name":"Gui-Tang Wang","email":"","orcid":"","institution":"Chinese Academy of Sciences","correspondingAuthor":false,"prefix":"","firstName":"Gui-Tang","middleName":"","lastName":"Wang","suffix":""},{"id":471998011,"identity":"45004c34-2fb4-41b2-bdff-56bce06d8929","order_by":10,"name":"Fei Liu","email":"","orcid":"","institution":"Xizang Autonomous Region Academy of Agricultural and Animal Husbandry Sciences","correspondingAuthor":false,"prefix":"","firstName":"Fei","middleName":"","lastName":"Liu","suffix":""},{"id":471998012,"identity":"18e75fa0-d997-4a98-b938-97c087c0e4e8","order_by":11,"name":"Benhe Zeng","email":"","orcid":"","institution":"Xizang Autonomous Region Academy of Agricultural and Animal Husbandry Sciences","correspondingAuthor":false,"prefix":"","firstName":"Benhe","middleName":"","lastName":"Zeng","suffix":""},{"id":471998013,"identity":"4b5ecd3e-083d-4267-abb3-e9cfab635829","order_by":12,"name":"Jinsong Chen","email":"","orcid":"","institution":"Sun Yat-Sen University","correspondingAuthor":false,"prefix":"","firstName":"Jinsong","middleName":"","lastName":"Chen","suffix":""}],"badges":[],"createdAt":"2025-05-18 23:05:10","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6693840/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6693840/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41467-026-70576-z","type":"published","date":"2026-03-12T04:00:00+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":107973385,"identity":"800a477b-e6bd-400f-9203-6fdc2658bad1","added_by":"auto","created_at":"2026-04-28 07:11:14","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":5644159,"visible":true,"origin":"","legend":"Article File","description":"","filename":"Manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6693840/v1_covered_61b5205f-ca32-40a4-b922-87af18d6d460.pdf"},{"id":84780674,"identity":"3bd74b11-bcd6-4680-b513-ab8886328ca0","added_by":"auto","created_at":"2025-06-17 09:26:59","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":22224610,"visible":true,"origin":"","legend":"Supplementary file","description":"","filename":"Supplementaryfile.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6693840/v1/bc27ec77b97de0aef0a1b172.pdf"},{"id":84781255,"identity":"3c25d80a-7144-44e5-9b4c-18eb7d10376d","added_by":"auto","created_at":"2025-06-17 09:34:59","extension":"xlsx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":2830580,"visible":true,"origin":"","legend":"Dataset 1","description":"","filename":"Supplementaryfile2.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-6693840/v1/b02518d8ed21f3df7afef3b7.xlsx"},{"id":84780666,"identity":"4736ffa3-a439-4d6e-a8e0-df310a72e338","added_by":"auto","created_at":"2025-06-17 09:26:59","extension":"pdf","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":161855,"visible":true,"origin":"","legend":"Reporting Summary","description":"","filename":"rs2.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6693840/v1/b52f982be150c6dfaa3d1613.pdf"}],"financialInterests":"There is \u003cb\u003eNO\u003c/b\u003e Competing Interest.","formattedTitle":"The evolution of mitogenomic architecture in Bilateria","fulltext":[],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":false,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":true,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":true,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"
[email protected]","identity":"nature-portfolio","isNatureJournal":true,"hasQc":false,"allowDirectSubmit":false,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"","title":"Nature Portfolio","twitterHandle":"","acdcEnabled":false,"dfaEnabled":false,"editorialSystem":"ejp","reportingPortfolio":"","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"mtDNA, mitochondrion, gene order, rearrangement, parasitism, metabolic rate, molecular evolution, purifying selection, locomotion, strand, animals ","lastPublishedDoi":"10.21203/rs.3.rs-6693840/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6693840/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"We used a dataset comprising over ten thousand bilaterian mitogenomes to address multiple open questions regarding the evolution of mitogenomic architecture. Double-stranded architecture is the most likely ancestral state for all major radiations (phylum and above), aside from Kinorhyncha, Syndermata and Brachiopoda. We found over twenty transitions from double-stranded to single-stranded (all genes on a single strand) mitogenomic architectures, as well as the first evidence of single-stranded to double-stranded transitions: one in Annelida and two or more in Mollusca. Gene order rearrangement rates (GORR) were strongly positively correlated with sequence evolution rates (r=0.69, R2≈35%). Both variables were increased in single-stranded mitogenomes, parasitic species, and those with low locomotory capacity. The latter two categories also exhibited increased proportions of single-stranded mitogenomes. Contrary to this, mitogenome size exhibited a negative correlation with both GORR and sequence evolutionary rates. Purifying selection plays a major role in the mitogenomic architectural evolution, but other variables need to be invoked to fully explain the observed patterns.","manuscriptTitle":"The evolution of mitogenomic architecture in Bilateria","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-17 09:26:54","doi":"10.21203/rs.3.rs-6693840/v1","editorialEvents":[],"status":"published","journal":{"display":true,"email":"
[email protected]","identity":"nature-communications","isNatureJournal":true,"hasQc":false,"allowDirectSubmit":false,"externalIdentity":"NCOMMS","sideBox":"Learn more about [Nature Communications](http://www.nature.com/ncomms/)","snPcode":"","submissionUrl":"https://mts-ncomms.nature.com/","title":"Nature Communications","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"ejp","reportingPortfolio":"Nature Communications","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"b8993ce7-68df-48de-aa3a-218aefc563e5","owner":[],"postedDate":"June 17th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":50123071,"name":"Biological sciences/Evolution/Molecular evolution"},{"id":50123072,"name":"Earth and environmental sciences/Ecology/Evolutionary ecology"}],"tags":[],"updatedAt":"2026-04-28T07:10:56+00:00","versionOfRecord":{"articleIdentity":"rs-6693840","link":"https://doi.org/10.1038/s41467-026-70576-z","journal":{"identity":"nature-communications","isVorOnly":false,"title":"Nature Communications"},"publishedOn":"2026-03-12 04:00:00","publishedOnDateReadable":"March 12th, 2026"},"versionCreatedAt":"2025-06-17 09:26:54","video":"","vorDoi":"10.1038/s41467-026-70576-z","vorDoiUrl":"https://doi.org/10.1038/s41467-026-70576-z","workflowStages":[]},"version":"v1","identity":"rs-6693840","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6693840","identity":"rs-6693840","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}
Text is read by the "Ask this paper" AI Q&A widget below.
Extraction quality varies by source — PMC NXML preserves structure
cleanly, OA-HTML may include some navigation residue, and OA-PDF can
have broken hyphenation. The publisher copy
(via DOI)
is the canonical version.