Fertility, mating behavior & group size - a unified empirical theory, hunter-gatherers to megacities

Fertility, mating behavior & group size - a unified empirical theory, hunter-gatherers to megacities | 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 Fertility, mating behavior & group size - a unified empirical theory, hunter-gatherers to megacities Robert Shuler This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7812686/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 As human groups expand, both fertility and mated ensemble size decline. Monogamy emerges in large societies as cooperation shifts from the shared outcomes of foraging bands to individuated, competitive exchange. This study develops a unified empirical framework linking fertility, mating behavior, and group size ( N ) across human evolution, from hunter-gatherer bands to modern megacities. Using ethnographic, archaeological, and demographic evidence, we define a composite measure of mating behavior ( M ) as mated ensemble size ( N e ) multiplied by normalized total fertility rate. The results formalize a feedback loop: higher fertility increases N , larger N suppresses fertility, and large- N systems become evolutionarily stable despite sub-replacement reproduction. Then we extend the framework to include the diffusion of polygenic traits that are beneficial to groups but deleterious to individuals . Using a minimal multilevel-selection model derived from the Price and the breeder’s equations, we evaluate how deme size modulates the fixation of such traits under Wright–Fisher drift and island-model exogamy. Results suggest that group-level benefits dominate within an intermediate band of N , producing an inverted-U relationship between group size and trait diffusion. Combined with the fertility– N feedback loop, this implies that modern large- N societies not only depress fertility but may also suppress the evolutionary maintenance of group-beneficial traits. Pronatalist policies may temporarily raise births but ultimately reinforce the fertility-suppressing dynamics of large- N . The findings raise the question of whether new cultural or institutional linkages can restore reproductive viability in complex societies. Biological sciences/Ecology Earth and environmental sciences/Ecology Biological sciences/Evolution Fertility decline mating systems group size evolutionary demography monogamy urbanization cultural evolution Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 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. 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-7812686","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":558816038,"identity":"166fe77a-1e00-4ea7-91c6-f192a808ae09","order_by":0,"name":"Robert 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0.001\u003c/em\u003e\u003c/p\u003e","description":"","filename":"Fig1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7812686/v1/9bb5ff6c8c6f717282486704.jpg"},{"id":98428921,"identity":"16478df9-573a-4777-a651-a2bedf1e3fd7","added_by":"auto","created_at":"2025-12-17 16:42:35","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":249550,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eMedian relatedness after 20 generations\u003c/strong\u003e as function of group size (\u003cem\u003eN\u003c/em\u003e), mated ensemble size (\u003cem\u003eM / 2\u003c/em\u003e), and exogamy rate.\u003c/p\u003e","description":"","filename":"Fig2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7812686/v1/4e4593e89b7e1ca62face7cd.jpg"},{"id":98059963,"identity":"96fe0d91-d100-4d23-9e5f-5b354a6e4a14","added_by":"auto","created_at":"2025-12-12 10:40:38","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":448942,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFertility-correlated factors by region. \u003c/strong\u003eBlue = spousal-age gap; gray = total fertility rate (\u003cem\u003eTFR\u003c/em\u003e); green = mean IQ scaled as (IQ − 75)/5. Shaded band marks replacement fertility. Cross-regional ordering reveals opposing gradients: higher intelligence aligns with narrower age gaps and sub-replacement fertility, while wider gaps correspond to \u003cem\u003eTFR\u003c/em\u003e \u0026gt; 3.5.\u003c/p\u003e","description":"","filename":"Fig3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7812686/v1/d59b8f5e08907c9d392751da.jpg"},{"id":98059970,"identity":"27b2a439-b893-4ec4-9caf-a5912deb1f49","added_by":"auto","created_at":"2025-12-12 10:40:38","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":511355,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003e(\u003c/strong\u003ea) US Birth Rate vs Household Income , (b) Fecundity vs Female Age, (c) Education \u0026amp; Age of 1\u003csup\u003est\u003c/sup\u003e Child by IQv(SMPY projection), (d) Mean Number of Children \u0026amp; Spacing by IQ (SMPY projection)\u003c/p\u003e","description":"","filename":"Figs4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7812686/v1/463bcaab0fa2da0664d3e90f.jpg"},{"id":98059971,"identity":"d193e0a4-7f50-47c4-b081-b3992b74801d","added_by":"auto","created_at":"2025-12-12 10:40:38","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":354610,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003e\u003cstrong\u003eDiffusion of Polygenic Group-beneficial Traits\u003c/strong\u003e\u003c/em\u003e\u003cem\u003e, (a) constant group fitness advantage and individual cost with diffusion, reversion to mean and exogamy 20%; (b) uses progressive trait fitness from Table 2.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"Figs5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7812686/v1/d14c63e4d808fac1be223083.jpg"},{"id":103506292,"identity":"52bfc06e-cfc8-46c4-b954-fab4d598205d","added_by":"auto","created_at":"2026-02-26 13:34:55","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2520684,"visible":true,"origin":"","legend":"","description":"","filename":"2025NHSSC1MatingBehaviorGroupSize.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7812686/v1_covered_27cd8e10-4312-45c2-985d-7cd952443603.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Fertility, mating behavior \u0026 group size - a unified empirical theory, hunter-gatherers to megacities","fulltext":[],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":false,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"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":"Fertility decline, mating systems, group size, evolutionary demography, monogamy, urbanization, cultural evolution","lastPublishedDoi":"10.21203/rs.3.rs-7812686/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7812686/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eAs human groups expand, both fertility and mated ensemble size decline. Monogamy emerges in large societies as cooperation shifts from the shared outcomes of foraging bands to individuated, competitive exchange. This study develops a unified empirical framework linking fertility, mating behavior, and group size (\u003cem\u003eN\u003c/em\u003e) across human evolution, from hunter-gatherer bands to modern megacities. Using ethnographic, archaeological, and demographic evidence, we define a composite measure of mating behavior (\u003cem\u003eM\u003c/em\u003e) as mated ensemble size (\u003cem\u003eN\u003c/em\u003e\u003csub\u003e\u003cem\u003ee\u003c/em\u003e\u003c/sub\u003e) multiplied by normalized total fertility rate. The results formalize a feedback loop: higher fertility increases \u003cem\u003eN\u003c/em\u003e, larger \u003cem\u003eN\u003c/em\u003e suppresses fertility, and large-\u003cem\u003eN\u003c/em\u003e systems become evolutionarily stable despite sub-replacement reproduction. Then we extend the framework to include the diffusion of polygenic traits that are \u003cem\u003ebeneficial to groups but deleterious to individuals\u003c/em\u003e. Using a minimal multilevel-selection model derived from the Price and the breeder\u0026rsquo;s equations, we evaluate how deme size modulates the fixation of such traits under Wright\u0026ndash;Fisher drift and island-model exogamy. Results suggest that group-level benefits dominate within an intermediate band of \u003cem\u003eN\u003c/em\u003e, producing an inverted-U relationship between group size and trait diffusion. Combined with the fertility\u0026ndash;\u003cem\u003eN\u003c/em\u003e feedback loop, this implies that modern large-\u003cem\u003eN\u003c/em\u003e societies not only depress fertility but may also suppress the evolutionary maintenance of group-beneficial traits. Pronatalist policies may temporarily raise births but ultimately reinforce the fertility-suppressing dynamics of large-\u003cem\u003eN\u003c/em\u003e. The findings raise the question of whether new cultural or institutional linkages can restore reproductive viability in complex societies.\u003c/p\u003e","manuscriptTitle":"Fertility, mating behavior \u0026amp; group size - a unified empirical theory, hunter-gatherers to megacities","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-12-12 10:40:33","doi":"10.21203/rs.3.rs-7812686/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","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}}],"origin":"","ownerIdentity":"32aa4905-46d6-4d56-85c2-ea936a4845ca","owner":[],"postedDate":"December 12th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":59488268,"name":"Biological sciences/Ecology"},{"id":59488269,"name":"Earth and environmental sciences/Ecology"},{"id":59488270,"name":"Biological sciences/Evolution"}],"tags":[],"updatedAt":"2026-02-23T20:24:27+00:00","versionOfRecord":[],"versionCreatedAt":"2025-12-12 10:40:33","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7812686","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7812686","identity":"rs-7812686","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}