Trade‑offs and timing of leaf defenses during development in co‑occurring evergreen and deciduous Quercus

Trade‑offs and timing of leaf defenses during development in co‑occurring evergreen and deciduous Quercus | 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 Trade‑offs and timing of leaf defenses during development in co‑occurring evergreen and deciduous Quercus Domingo Sancho-Knapik, Nerea Urcola-Lázaro, David Alonso‑Forn, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8639643/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 9 You are reading this latest preprint version Abstract Context: Young, expanding leaves are nutrient-rich and mechanically weak, making them highly attractive to herbivores. How defense timing and allocation differ during development between co-occurring evergreen and deciduous species remains unclear. Aims: We tested whether evergreen oaks invest more in direct defenses during leaf development than deciduous oaks, and whether each habit prioritizes distinct defensive traits across stages. Methods: In a Mediterranean common garden, we monitored co-occurring evergreen Quercus rotundifolia and deciduous Q. faginea from budburst to leaf maturity, measuring growth and defense traits. Seasonal trajectories and toughness–tannin relationships were analyzed, and patterns validated with three additional evergreen and three deciduous congeners. Results: Leaf area reached its maximum before LMA, creating a juvenile phase with low fibre content and toughness. The evergreen oak increased condensed tannins concentration early, then work of fracture; the deciduous oak grew faster initially, then reinforced mechanically and delayed tannins. Across stages, the evergreen kept higher work of fracture and condensed tannins, while the deciduous prioritized early growth; defense trade-off patterns were habit-specific and consistent across species. Conclusion: Leaf-habit–specific defense schedules shape protection: evergreens front-load chemical defenses before structural hardening, while deciduous oaks prioritize tolerance and earlier mechanical reinforcement, patterns with potential implications for herbivore pressure and associational dynamics in mixed stands. Condensed tannins defense trade‑offs leaf ontogeny relative growth rate work of fracture Full Text Additional Declarations No competing interests reported. Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Revision requested 02 Apr, 2026 Reviews received at journal 19 Mar, 2026 Reviews received at journal 09 Mar, 2026 Reviewers agreed at journal 04 Mar, 2026 Reviewers agreed at journal 18 Feb, 2026 Reviewers invited by journal 17 Feb, 2026 Editor assigned by journal 17 Feb, 2026 Submission checks completed at journal 16 Feb, 2026 First submitted to journal 19 Jan, 2026 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. 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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-8639643","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":593564662,"identity":"4ad032d6-e3ab-4ba5-89df-d9449acd0a18","order_by":0,"name":"Domingo Sancho-Knapik","email":"","orcid":"","institution":"Centro de Investigación y Tecnología Agroalimentaria de Aragón","correspondingAuthor":false,"prefix":"","firstName":"Domingo","middleName":"","lastName":"Sancho-Knapik","suffix":""},{"id":593564663,"identity":"a8558ace-aa9d-44dc-aa46-8bbc5ca42695","order_by":1,"name":"Nerea 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How defense timing and allocation differ during development between co-occurring evergreen and deciduous species remains unclear.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAims: \u003c/strong\u003eWe tested whether evergreen oaks invest more in direct defenses during leaf development than deciduous oaks, and whether each habit prioritizes distinct defensive traits across stages.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMethods: \u003c/strong\u003eIn a Mediterranean common garden, we monitored co-occurring evergreen \u003cem\u003eQuercus rotundifolia\u003c/em\u003eand deciduous \u003cem\u003eQ. faginea\u003c/em\u003e from budburst to leaf maturity, measuring growth and defense traits. Seasonal trajectories and toughness–tannin relationships were analyzed, and patterns validated with three additional evergreen and three deciduous congeners.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eResults: \u003c/strong\u003eLeaf area reached its maximum before LMA, creating a juvenile phase with low fibre content and toughness. The evergreen oak increased condensed tannins concentration early, then work of fracture; the deciduous oak grew faster initially, then reinforced mechanically and delayed tannins. Across stages, the evergreen kept higher work of fracture and condensed tannins, while the deciduous prioritized early growth; defense trade-off patterns were habit-specific and consistent across species.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConclusion: \u003c/strong\u003eLeaf-habit–specific defense schedules shape protection: evergreens front-load chemical defenses before structural hardening, while deciduous oaks prioritize tolerance and earlier mechanical reinforcement, patterns with potential implications for herbivore pressure and associational dynamics in mixed stands.\u003c/p\u003e","manuscriptTitle":"Trade‑offs and timing of leaf defenses during development in co‑occurring evergreen and deciduous Quercus","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-02-19 12:20:02","doi":"10.21203/rs.3.rs-8639643/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-04-02T08:12:47+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-19T10:31:42+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-09T18:36:53+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"263442645157834236307177926649827190238","date":"2026-03-04T19:40:00+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"237222981520345086259066934295444326397","date":"2026-02-18T17:21:53+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-02-17T10:22:30+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-02-17T09:25:29+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-02-17T04:54:07+00:00","index":"","fulltext":""},{"type":"submitted","content":"Annals of Forest Science","date":"2026-01-19T12:33:21+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"annals-of-forest-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"","sideBox":"Learn more about [Annals of Forest Science](https://link.springer.com/journal/13595)","snPcode":"13595","submissionUrl":"https://submission.springernature.com/new-submission/13595/3","title":"Annals of Forest Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"111518c5-770a-4931-9ca9-f8687bdaca72","owner":[],"postedDate":"February 19th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-04-24T13:54:50+00:00","versionOfRecord":[],"versionCreatedAt":"2026-02-19 12:20:02","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8639643","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8639643","identity":"rs-8639643","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}