Modelling the long-term effects of fire frequency, intensity and rainfall variability on tree-grass coexistence in a semi-arid savanna, Namibia | 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 Modelling the long-term effects of fire frequency, intensity and rainfall variability on tree-grass coexistence in a semi-arid savanna, Namibia Jesaya Nakanyala, Johaness Amutenya This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7585321/v1 This work is licensed under a CC BY 4.0 License Status: Under Revision Version 1 posted 11 You are reading this latest preprint version Abstract Background: The semi-arid savannas of Namibia are shaped by complex interactions among rainfall variability, fire regimes, and vegetation dynamics. However, the long-term consequences of these drivers for tree and grass coexistence remain poorly understood. We developed a simulation model in R to investigate the effects of fire frequency and intensity on grass and tree biomass in semi-arid Namibian savannas. The model incorporates rainfall variability, logistic vegetation growth, and stochastic fire events, capturing interactions between fuel availability, fire probability, and vegetation responses. Results: Grass biomass responded rapidly to wet-season rainfall pulses and showed resilience to fire, whereas tree biomass accumulated more slowly and was strongly suppressed by frequent, high-intensity fires. Low-frequency, mild fires promoted tree establishment and coexistence with grasses, while high-frequency, severe fires maintained grass-dominated states. Intermediate fire regimes fostered dynamic equilibria, with both vegetation types oscillating in response to rainfall and disturbance. Interannual rainfall variability amplified biomass fluctuations, occasionally enhancing tree recruitment in wet years and suppressing it in dry periods, underscoring the role of stochastic drivers in semi-arid ecosystems. Conclusion: Our results emphasize that adaptive fire management is crucial for sustaining productive and resilient savannas. Conducting controlled burns every 3 to 5 years helps reduce fuel loads, prevents excessive accumulation of grass biomass that can drive severe wildfires, and creates opportunities for tree recruitment. Ideally, burns should occur early in the dry season, when fuel moisture is higher, to limit fire intensity and reduce tree mortality. Late dry season fires should be minimized, as they tend to be hotter and more destructive. By integrating these practices with rainfall monitoring, for example scheduling burns after years of high grass growth, rangeland managers can balance forage availability, mitigate wildfire risk, and support the coexistence of trees and grasses in Namibia’s semi-arid savannas. Fire tree grass savanna simulation Full Text Additional Declarations No competing interests reported. Cite Share Download PDF Status: Under Revision Version 1 posted Editorial decision: Revision requested 17 Dec, 2025 Reviews received at journal 16 Dec, 2025 Reviews received at journal 11 Dec, 2025 Reviewers agreed at journal 26 Nov, 2025 Reviewers agreed at journal 26 Nov, 2025 Reviews received at journal 19 Nov, 2025 Reviewers agreed at journal 09 Oct, 2025 Reviewers invited by journal 06 Oct, 2025 Editor assigned by journal 03 Oct, 2025 Submission checks completed at journal 03 Oct, 2025 First submitted to journal 02 Oct, 2025 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-7585321","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":529327978,"identity":"8c139d37-afe2-46ae-ab91-dc01132d937d","order_by":0,"name":"Jesaya Nakanyala","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA5klEQVRIiWNgGAWjYBACxgYGZjCDj4H5AJCSkCFeCxsDWwJICw8xFsG08BiAaMJamGc3Pzb4ueOwPBv7mc+vbtRY8DCwHz66Aa/D5hwzTuw9c9iwjSd3m3XOMaDDeNLSbuDVMiPB+ABv22HGNobcbcY5bEAtEjxmBLSkfz74t+2wfRv/m2fGOf+I0pJjnAy0JbFNIof5cW4bcVqKjWXb0pPbJJ6ZMef2SfCwEfKL4Yz0zZJv26xt+/mTH3/O+VYnx89++Bh+LQ1gqhlEsEmASXzKQUAeQtWBCOYPhFSPglEwCkbByAQA2C5GR3Ry0uIAAAAASUVORK5CYII=","orcid":"","institution":"University of Namibia","correspondingAuthor":true,"prefix":"","firstName":"Jesaya","middleName":"","lastName":"Nakanyala","suffix":""},{"id":529327979,"identity":"c9f9f370-8361-48c3-93ef-08dddd342240","order_by":1,"name":"Johaness Amutenya","email":"","orcid":"","institution":"University of Namibia","correspondingAuthor":false,"prefix":"","firstName":"Johaness","middleName":"","lastName":"Amutenya","suffix":""}],"badges":[],"createdAt":"2025-09-10 17:53:09","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7585321/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7585321/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":93714010,"identity":"81335ef3-61fc-439c-bf15-fb8d18096430","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"json","order_by":0,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":5107,"visible":true,"origin":"","legend":"","description":"","filename":"37c7bac5f0c74596b7fbc3a69799ec51.json","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/983d9060f46c72a4c5c34db0.json"},{"id":93714012,"identity":"a838099f-7519-421c-8c0b-213714bbebac","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"xml","order_by":1,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":63888,"visible":true,"origin":"","legend":"","description":"","filename":"37c7bac5f0c74596b7fbc3a69799ec511enriched.xml","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/91e861d9a501a19add738625.xml"},{"id":93714334,"identity":"a404104f-4594-4898-9730-bc83c8fdf113","added_by":"auto","created_at":"2025-10-16 19:03:08","extension":"pdf","order_by":2,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":36190,"visible":true,"origin":"","legend":"","description":"","filename":"Coverletter.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/78ef1a35a7b132bf84de2fea.pdf"},{"id":93714011,"identity":"25758bf5-7d12-4641-8c54-22208b23de37","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"png","order_by":3,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1099861,"visible":true,"origin":"","legend":"","description":"","filename":"Fig.1.png","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/eb3777ceffb4296801d96063.png"},{"id":93714008,"identity":"678efb77-a7b2-49bb-9179-833c1f667799","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"png","order_by":4,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":84249,"visible":true,"origin":"","legend":"","description":"","filename":"Fig.2.png","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/af60b4b4916e50723c4e5354.png"},{"id":93714015,"identity":"97dbd2a3-a763-43d4-bb15-dc336e6d4c2a","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"pdf","order_by":5,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1546309,"visible":true,"origin":"","legend":"","description":"","filename":"ModellingthelongtermeffectsoffirefrequencyintensityandrainfallvariabilityontreegrasscoexistenceinasemiaridsavannaNamibia.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/7f83f03a443c9467067395bd.pdf"},{"id":93714014,"identity":"409547d9-f82c-413c-80a1-75dfcdf0248f","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"pdf","order_by":6,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":48058,"visible":true,"origin":"","legend":"","description":"","filename":"Responsetoeditorscomments.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/efbdcd71b27afcdfc7f385fc.pdf"},{"id":93714335,"identity":"27e7637d-c038-4f98-85e5-802031eebf76","added_by":"auto","created_at":"2025-10-16 19:03:08","extension":"eps","order_by":7,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":2890,"visible":true,"origin":"","legend":"","description":"","filename":"empty.eps","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/a558e9324abded0a0f602e86.eps"},{"id":93714848,"identity":"7e0b6e67-34c8-4582-b995-49da5eede54b","added_by":"auto","created_at":"2025-10-16 19:11:08","extension":"eps","order_by":8,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":91593,"visible":true,"origin":"","legend":"","description":"","filename":"fig.eps","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/73334bb3e6de9c39605a539c.eps"},{"id":93714344,"identity":"31a4122e-cc91-4ee8-84e1-d731a9558c65","added_by":"auto","created_at":"2025-10-16 19:03:08","extension":"bst","order_by":9,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":146013,"visible":true,"origin":"","legend":"","description":"","filename":"snapacite.bst","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/9a4958cc64e3cc126dca9edc.bst"},{"id":93714337,"identity":"2b821e3b-dc6a-420a-b693-253fdaaae611","added_by":"auto","created_at":"2025-10-16 19:03:08","extension":"bst","order_by":10,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":29828,"visible":true,"origin":"","legend":"","description":"","filename":"snaps.bst","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/35357e1f3dd01121008c469f.bst"},{"id":93714850,"identity":"76f30cbb-d8b9-4d4d-b674-6ce9ef09924f","added_by":"auto","created_at":"2025-10-16 19:11:08","extension":"pdf","order_by":11,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":421391,"visible":true,"origin":"","legend":"","description":"","filename":"snarticle.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/4e5e0f55d16a6d9b27024369.pdf"},{"id":93714024,"identity":"b2e7a7e1-e36f-4642-989a-a022dfd54bea","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"bst","order_by":12,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":35515,"visible":true,"origin":"","legend":"","description":"","filename":"snbasic.bst","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/5efad6b9a2cc7faae19f5415.bst"},{"id":93714017,"identity":"c75773bd-fa90-4063-aa71-c6aae7106c40","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"bst","order_by":13,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":33968,"visible":true,"origin":"","legend":"","description":"","filename":"snchicago.bst","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/67ff29b4d2dded1cdd3318e4.bst"},{"id":93714336,"identity":"04bcd7e7-da72-4be9-b7c8-ce5540c36110","added_by":"auto","created_at":"2025-10-16 19:03:08","extension":"cls","order_by":14,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":55857,"visible":true,"origin":"","legend":"","description":"","filename":"snjnl.cls","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/73d3b6e6c8d8a9beb9d4180f.cls"},{"id":93714026,"identity":"7f0c133e-081f-492b-a917-6a0e90351740","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"bst","order_by":15,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":64023,"visible":true,"origin":"","legend":"","description":"","filename":"snmathphysay.bst","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/0b4d4d073ccb184231e03a26.bst"},{"id":93714849,"identity":"7f7dd7d2-702a-4ead-9273-39988f1947cd","added_by":"auto","created_at":"2025-10-16 19:11:08","extension":"bst","order_by":16,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":64166,"visible":true,"origin":"","legend":"","description":"","filename":"snmathphysnum.bst","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/9aeb4d919d1822142cb309b1.bst"},{"id":93714343,"identity":"3ba539dc-e840-4bdc-ad8f-dd73ba3b8e22","added_by":"auto","created_at":"2025-10-16 19:03:08","extension":"bst","order_by":17,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":37333,"visible":true,"origin":"","legend":"","description":"","filename":"snnature.bst","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/fb684f4068f3a2e2bcd3d32f.bst"},{"id":93714028,"identity":"064db427-d055-46c1-b0fc-22d2bc81a944","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"bst","order_by":18,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":39951,"visible":true,"origin":"","legend":"","description":"","filename":"snvancouveray.bst","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/779b6b176aa86d7e42fdfbc7.bst"},{"id":93714341,"identity":"d932e9ee-6ca1-4ad9-8361-599d84c8232b","added_by":"auto","created_at":"2025-10-16 19:03:08","extension":"bst","order_by":19,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":40758,"visible":true,"origin":"","legend":"","description":"","filename":"snvancouvernum.bst","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/0aa5fa051d864f2ca079b951.bst"},{"id":93714021,"identity":"f019240c-e5a1-4552-b001-766937c79533","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"pdf","order_by":20,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":418495,"visible":true,"origin":"","legend":"","description":"","filename":"usermanual.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/8778546c002e5855f66326ae.pdf"},{"id":93714339,"identity":"5410dc00-c12d-4fca-b214-e7746141be81","added_by":"auto","created_at":"2025-10-16 19:03:08","extension":"xml","order_by":23,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":66353,"visible":true,"origin":"","legend":"","description":"","filename":"37c7bac5f0c74596b7fbc3a69799ec511structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/fb12ecb55a649ee77f8f314c.xml"},{"id":93714030,"identity":"f6526828-d16d-43ca-916c-41ffe480e3bb","added_by":"auto","created_at":"2025-10-16 18:55:08","extension":"html","order_by":24,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":75784,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1/ea216e39146678424e0f0430.html"},{"id":93714985,"identity":"dc68c5aa-9521-42de-b346-ac8992e07744","added_by":"auto","created_at":"2025-10-16 19:19:10","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1843872,"visible":true,"origin":"","legend":"","description":"","filename":"ModellingthelongtermeffectsoffirefrequencyintensityandrainfallvariabilityontreegrasscoexistenceinasemiaridsavannaNamibia.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7585321/v1_covered_738c398c-5629-4d67-af48-d4713931094b.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Modelling the long-term effects of fire frequency, intensity and rainfall variability on tree-grass coexistence in a semi-arid savanna, Namibia ","fulltext":[],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":false,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"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":"fire-ecology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"feco","sideBox":"Learn more about [Fire Ecology](https://www.springer.com/journal/42408)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/feco/default.aspx","title":"Fire Ecology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Fire, tree, grass, savanna, simulation","lastPublishedDoi":"10.21203/rs.3.rs-7585321/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7585321/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cstrong\u003eBackground: \u003c/strong\u003eThe semi-arid savannas of Namibia are shaped by complex interactions among rainfall variability, fire regimes, and vegetation dynamics. However, the long-term consequences of these drivers for tree and grass coexistence remain poorly understood. We developed a simulation model in R to investigate the effects of fire frequency and intensity on grass and tree biomass in semi-arid Namibian savannas. The model incorporates rainfall variability, logistic vegetation growth, and stochastic fire events, capturing interactions between fuel availability, fire probability, and vegetation responses.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eResults: \u003c/strong\u003eGrass biomass responded rapidly to wet-season rainfall pulses and showed resilience to fire, whereas tree biomass accumulated more slowly and was strongly suppressed by frequent, high-intensity fires. Low-frequency, mild fires promoted tree establishment and coexistence with grasses, while high-frequency, severe fires maintained grass-dominated states. Intermediate fire regimes fostered dynamic equilibria, with both vegetation types oscillating in response to rainfall and disturbance. Interannual rainfall variability amplified biomass fluctuations, occasionally enhancing tree recruitment in wet years and suppressing it in dry periods, underscoring the role of stochastic drivers in semi-arid ecosystems.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConclusion:\u003c/strong\u003e Our results emphasize that adaptive fire management is crucial for sustaining productive and resilient savannas. Conducting controlled burns every 3 to 5 years helps reduce fuel loads, prevents excessive accumulation of grass biomass that can drive severe wildfires, and creates opportunities for tree recruitment. Ideally, burns should occur early in the dry season, when fuel moisture is higher, to limit fire intensity and reduce tree mortality. Late dry season fires should be minimized, as they tend to be hotter and more destructive. By integrating these practices with rainfall monitoring, for example scheduling burns after years of high grass growth, rangeland managers can balance forage availability, mitigate wildfire risk, and support the coexistence of trees and grasses in Namibia’s semi-arid savannas.\u003c/p\u003e","manuscriptTitle":"Modelling the long-term effects of fire frequency, intensity and rainfall variability on tree-grass coexistence in a semi-arid savanna, Namibia","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-16 18:55:03","doi":"10.21203/rs.3.rs-7585321/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-12-17T10:46:02+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-12-16T12:54:53+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-12-11T12:44:43+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"114629040404219446769811353390624199040","date":"2025-11-26T12:29:12+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"232800270638318927518075361734164896631","date":"2025-11-26T11:11:17+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-11-19T16:15:43+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"243871654358038450021077697264982470116","date":"2025-10-09T22:05:31+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-10-06T08:29:54+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-10-03T13:17:32+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-10-03T12:13:15+00:00","index":"","fulltext":""},{"type":"submitted","content":"Fire Ecology","date":"2025-10-02T19:20:34+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"
[email protected]","identity":"fire-ecology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"feco","sideBox":"Learn more about [Fire Ecology](https://www.springer.com/journal/42408)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/feco/default.aspx","title":"Fire Ecology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"6f53aad5-c67b-40dd-bcd7-7b8e365517f0","owner":[],"postedDate":"October 16th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"in-revision","subjectAreas":[],"tags":[],"updatedAt":"2025-12-17T10:53:45+00:00","versionOfRecord":[],"versionCreatedAt":"2025-10-16 18:55:03","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7585321","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7585321","identity":"rs-7585321","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.