Morphological characterization of anterior axial bifurcation in Holothuria (Halodeima) grisea and other puzzling occurrences in Holothuroidea | 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 Research Article Morphological characterization of anterior axial bifurcation in Holothuria (Halodeima) grisea and other puzzling occurrences in Holothuroidea Guilherme Sabino Rupp, Luciana Martins, Camilla Souto, Jean-François Hamel, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4145043/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract The present study explores rare but taxonomically and geographically broad cases of anterior body axial bifurcation in adult sea cucumbers (Echinodermata: Holothuroidea). The case of Holothuria (Halodeima) grisea, a common intertidal species along the Brazilian coast, was studied externally and internally. Individuals of Apostichopus californicus in the United States of America, H. (Mertensiothuria) leucospilota in La Réunion (France), H. (Panningothuria) forskali in Spain, and Parastichopus tremulus in Norway, were also reported to display similar anterior body axial bifurcation. In each of them, the anterior body presented two mouths with their respective crown of tentacles. In vivo observations of H. (Halodeima) grisea showed that the two tentacle crowns and mouths were independently functional. Micro-computed tomography (μCT) revealed that the anterior region of the body (up to the pharynx) was duplicated. Internally, two aquapharyngeal bulbs were found, each with their own calcareous ring (one with two plates missing), madreporite, and Polian vesicles. The body presented a normal set of five longitudinal muscle bands, some of which bifurcated orally to support more than one calcareous ring plate. This is the first report of such abnormalities in Holothuroidea and hypotheses that may explain such phenomenon are discussed. Evolutionary Biology Developmental Biology Taxonomy Abnormal morphology holothurian sea cucumber axial bifurcation microcomputed tomography Holothuria Full Text Additional Declarations The authors declare no competing interests. Supporting Information S2 is not available with this version. Cite Share Download PDF Status: Posted 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. 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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-4145043","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":282370038,"identity":"e246ec6f-34e3-41c0-ae24-abc1b131c9cb","order_by":0,"name":"Guilherme Sabino Rupp","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA5UlEQVRIiWNgGAWjYLCCD2CSsQHGZyOog3EGyVqYedAE8Gsxn5F77LFNxR15+ejDDYw/ahjs+iUS2B5X4NEicyMv3TjnzDPDjecSG5h5jjEkz5yRwG54Bo8WCYkcM+nctsOMG3sYG5iB3kk2OHOATbKBkBbLf4ftQVoYfxKthbHhcOJ8HqAVvA0MdgbHGwho4XmXbthz7FnyBqCWwzzHJBIk2xvbDfFqYc899uBHzR3b+T3sDx/+qLGx52dmPvYQnxYGBh5QLBxgMDgAIhkkEhuQ4hS/FnmoOnsCykfBKBgFo2AEAgBC1UpigvJhKgAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0002-5476-9689","institution":"e Institution for Agricultural Research and Rural Extension, Epagri","correspondingAuthor":true,"prefix":"","firstName":"Guilherme","middleName":"Sabino","lastName":"Rupp","suffix":""},{"id":282370039,"identity":"47a623bd-9bfe-48e9-b023-103a4fec683b","order_by":1,"name":"Luciana Martins","email":"","orcid":"https://orcid.org/0000-0002-8107-3265","institution":"Oceanographic Institute, University of São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Luciana","middleName":"","lastName":"Martins","suffix":""},{"id":282370040,"identity":"b82b8ac2-2dce-466a-9a69-a255382b21fd","order_by":2,"name":"Camilla Souto","email":"","orcid":"https://orcid.org/0000-0001-5575-8263","institution":"School of Natural Sciences \u0026 Mathematics, USAStockton University, Galloway,","correspondingAuthor":false,"prefix":"","firstName":"Camilla","middleName":"","lastName":"Souto","suffix":""},{"id":282370041,"identity":"813ef17e-132f-4b8d-971a-0494e4d447ac","order_by":3,"name":"Jean-François Hamel","email":"","orcid":"https://orcid.org/0000-0002-0435-4127","institution":"Society for the Exploration and Valuing of the Environment (SEVE), Portugal Cove-St","correspondingAuthor":false,"prefix":"","firstName":"Jean-François","middleName":"","lastName":"Hamel","suffix":""},{"id":282370042,"identity":"b659316e-6148-4356-9ecd-95ddc0b9b570","order_by":4,"name":"Annie Mercier","email":"","orcid":"https://orcid.org/0000-0003-0807-2378","institution":"Department of Ocean Sciences, Memorial University, St. John’s, NL, A1C 5S7, Canada","correspondingAuthor":false,"prefix":"","firstName":"Annie","middleName":"","lastName":"Mercier","suffix":""}],"badges":[],"createdAt":"2024-03-21 17:02:41","currentVersionCode":1,"declarations":{"humanSubjects":false,"vertebrateSubjects":false,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":false},"doi":"10.21203/rs.3.rs-4145043/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4145043/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":53263028,"identity":"fe5d21fe-7bdb-4918-83ed-e52b6b785f36","added_by":"auto","created_at":"2024-03-22 15:00:01","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1247062,"visible":true,"origin":"","legend":"","description":"","filename":"Ruppetal.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4145043/v1_covered_603da66a-047a-460d-ab0a-533ca7c96059.pdf"}],"financialInterests":"\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\n\u003cp\u003eSupporting Information S2 is not available with this version.\u003c/p\u003e","formattedTitle":"\u003cp\u003eMorphological characterization of anterior axial bifurcation in Holothuria (Halodeima) grisea and other puzzling occurrences in Holothuroidea\u003c/p\u003e","fulltext":[],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":false,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":true,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":true,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"
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