Walking separately, together, or in mixed groups: Energetic modeling of hominin group locomotion using the Engare Sero archaeological footprint assemblage

Walking separately, together, or in mixed groups: Energetic modeling of hominin group locomotion using the Engare Sero archaeological footprint assemblage | 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 Walking separately, together, or in mixed groups: Energetic modeling of hominin group locomotion using the Engare Sero archaeological footprint assemblage Guillermo Zorrilla-Revilla This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9180888/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 Locomotor coordination is a fundamental pillar of human sociality, yet reconstructing group dynamics in deep time remains a significant challenge. The Late Pleistocene footprint site of Engare Sero (Tanzania) provides a unique record of anatomically modern humans moving collectively across a volcanic landscape. While previous studies have inferred group composition and speed from footprint morphology, the energetic implications of these movement patterns have remained unexplored. This study models the Cost of Transport (CoT) and energetic penalties for three representative profiles: adult male, adult female, and non-adult, using experimentally validated quadratic equations. Multiple walking scenarios derived from the Engare Sero assemblage were evaluated, including age-sex-specific means, directional mean speeds (northeast [NE] and southwest [SW]), and the total group average, which were compared with the calculated Optimal Locomotion Speed (OLS) for each profile. The results identify an energetically efficient range between 1.35 and 1.41 m·s⁻¹ (4.86–5.08 km·h⁻¹), where individual optima overlap. The reconstructed group mean speed (1.41 m·s⁻¹) yielded the mathematically lowest overall penalty (0.26%). However, given the female-biased demography of the site (77% adult females), the SW scenario (1.35 m·s⁻¹) appears demographically favorable, reducing the energetic penalty for the majority group to 0.09%. These findings support the interpretation of cooperative mobility in the Late Pleistocene and suggest that prehistoric human locomotion balanced metabolic efficiency with group cohesion. Hominin footprints human behavioral ecology energy expenditure hunter-gatherers Engare Sero group locomotion Full Text Additional Declarations No competing interests reported. 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. 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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-9180888","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":610306150,"identity":"2d80dd8a-252b-4f77-9697-d6f7fa0fa548","order_by":0,"name":"Guillermo Zorrilla-Revilla","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA7ElEQVRIiWNgGAWjYFACxsYDDxA8G4gIAS0NBxIQvDSICCF7kLUchojgUy4fkQy0peJeND//4WOfKyrO261tPwy0pcYmGpcWwxuJQC1ninNnNhxLnnnmzO3kbWeAIgzH0nIbcGmZAVSQ2JaQu+FgjzFjY9vtZLMDQBHGhsMEtPwDajnMA9Ty71yy2fmH+LXIS4C0NAC1HANpaThgZ3aDgC0GPEAzE44l5M7sYUtmBPonwewGWAS3X+Tb0x8++FCTkNvPf/gwY0ONnb3ZebCIDW5bDqAJJIJVJuBQDrYF3Sx7PIpHwSgYBaNghAIAThptCAD8SPsAAAAASUVORK5CYII=","orcid":"","institution":"University of Burgos","correspondingAuthor":true,"prefix":"","firstName":"Guillermo","middleName":"","lastName":"Zorrilla-Revilla","suffix":""}],"badges":[],"createdAt":"2026-03-20 16:53:31","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9180888/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9180888/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":106078204,"identity":"aa33bf69-8844-4ff1-a1ab-242c6f09425d","added_by":"auto","created_at":"2026-04-03 07:57:51","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":723540,"visible":true,"origin":"","legend":"","description":"","filename":"TextPageAAS.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9180888/v1_covered_d14b5bd3-8c8e-4c50-b47d-7f0cc786993c.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Walking separately, together, or in mixed groups: Energetic modeling of hominin group locomotion using the Engare Sero archaeological footprint assemblage","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":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Hominin footprints, human behavioral ecology, energy expenditure, hunter-gatherers, Engare Sero, group locomotion","lastPublishedDoi":"10.21203/rs.3.rs-9180888/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9180888/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Locomotor coordination is a fundamental pillar of human sociality, yet reconstructing group dynamics in deep time remains a significant challenge. 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