Quantifying the trade-off between spring phenology and lethal frost risk: a global meta-analysis

Quantifying the trade-off between spring phenology and lethal frost risk: a global meta-analysis | 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 Quantifying the trade-off between spring phenology and lethal frost risk: a global meta-analysis Jin-Sheng He, zhengjie yan, cheng chen, yue liu, Yujiang Li, Huiying Liu, and 7 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6668207/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 05 Mar, 2026 Read the published version in Nature Communications → Version 1 posted You are reading this latest preprint version Abstract The timing of spring phenology reflects a critical trade-off between growing season extension and frost risk avoidance, the balance of which governs plant fitness and shapes plant species’ distributions. However, few quantitative assessments of the spring lethal frost risk exist, meaning that this trade-off is largely untested empirically. Here, we present the first global meta-analysis encompassing 193 plant species across 126 study sites, finding consistently high freezing resistance (to -12°C) with remarkably low lethal frost risk (safety margin of 17°C) during spring emergence. Under projected climate warming scenarios, predicted advances in spring phenology did not affect the lethal frost risk for two plausible warming scenarios (SSP1-2.6 and SSP2-4.5), this was likely due to reduced temperature sensitivity, consistent with trade-off predictions. By contrast, in the high-warming scenario (SSP5-8.5), the freezing safety margin was predicted to expand (by ~ 1°C), resulting in a lower lethal frost risk, this may compensate for the diminished freezing resistance predicted under warming. Our findings challenge the conventional view of simple-environmental factors moderate spring phenology and highlight the necessity of incorporating biotic factors (e.g., freezing resistance) and biotic processes (e.g., early growth versus frost risk trade-offs) into next-generation phenological models to enhance prediction accuracy under climate change. Earth and environmental sciences/Ecology/Climate-change ecology/Phenology Earth and environmental sciences/Ecology/Evolutionary ecology Full Text Additional Declarations There is NO Competing Interest. Supplementary Files appendix.docx Supporting information for: Quantifying the trade-off between spring phenology and lethal frost risk: a global meta-analysis rs.pdf Reporting Summary Cite Share Download PDF Status: Published Journal Publication published 05 Mar, 2026 Read the published version in Nature Communications → 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-6668207","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":469322145,"identity":"516fd5d2-e59d-45bf-93c2-d8b398bb50ad","order_by":0,"name":"Jin-Sheng He","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAu0lEQVRIiWNgGAWjYBACxgYeBoYPDAkgtgHxWhhnkKSFgYGHgZmHJC3M/WcPPrbdkZbYwN68TYKh5g4RDpuRl2yceyYnsYHnWJkEw7FnxGjhMZPObatIbJDIMZNgbDhMhJb+M+a/LUFa5N8Qq6Uhx4yZsQ3oMAkeYrXMyDGW7D2TZtzGk1ZskXCMCC2G/WcMP/zckSzbz354440PNcRoaQC7joGBDcRLIKyBgUGeAaplFIyCUTAKRgFOAABr7jfOjaFe9wAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0001-5081-3569","institution":"Lanzhou University","correspondingAuthor":true,"prefix":"","firstName":"Jin-Sheng","middleName":"","lastName":"He","suffix":""},{"id":469322146,"identity":"ce3f0157-34b3-4b8c-b1e2-b6a621060aa3","order_by":1,"name":"zhengjie yan","email":"","orcid":"","institution":"State Key Laboratory of Herbage Improvement and Grassland Agro-Ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou, China","correspondingAuthor":false,"prefix":"","firstName":"zhengjie","middleName":"","lastName":"yan","suffix":""},{"id":469322147,"identity":"cbcc5172-f0e6-4df2-8df0-c624db4a6e8a","order_by":2,"name":"cheng chen","email":"","orcid":"","institution":"State Key Laboratory of Herbage Improvement and Grassland Agro-Ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou, China","correspondingAuthor":false,"prefix":"","firstName":"cheng","middleName":"","lastName":"chen","suffix":""},{"id":469322148,"identity":"41ba7372-6e8b-4f63-b096-fb59aa4a8dc1","order_by":3,"name":"yue liu","email":"","orcid":"","institution":"State Key Laboratory of Herbage Improvement and Grassland Agro-Ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou, China","correspondingAuthor":false,"prefix":"","firstName":"yue","middleName":"","lastName":"liu","suffix":""},{"id":469322149,"identity":"700af091-f1a8-4aeb-923c-72befe60a3ad","order_by":4,"name":"Yujiang Li","email":"","orcid":"https://orcid.org/0009-0007-8941-0477","institution":"Lanzhou University","correspondingAuthor":false,"prefix":"","firstName":"Yujiang","middleName":"","lastName":"Li","suffix":""},{"id":469322150,"identity":"69970729-e408-4c56-84c7-22a4f76b96a2","order_by":5,"name":"Huiying Liu","email":"","orcid":"https://orcid.org/0000-0001-8903-6103","institution":"East China Normal University","correspondingAuthor":false,"prefix":"","firstName":"Huiying","middleName":"","lastName":"Liu","suffix":""},{"id":469322151,"identity":"af66f8b6-2a1d-4d8c-b785-93252b37b8aa","order_by":6,"name":"Hao Wang","email":"","orcid":"https://orcid.org/0000-0001-9115-1290","institution":"Lanzhou University","correspondingAuthor":false,"prefix":"","firstName":"Hao","middleName":"","lastName":"Wang","suffix":""},{"id":469322152,"identity":"4a0035b9-97da-4cd7-afd6-8a74a3a6bc4e","order_by":7,"name":"Tao Yan","email":"","orcid":"","institution":"State Key Laboratory of Herbage Improvement and Grassland Agro-Ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou, China","correspondingAuthor":false,"prefix":"","firstName":"Tao","middleName":"","lastName":"Yan","suffix":""},{"id":469322153,"identity":"bc4fd7db-c780-4727-8e06-42e8be654b0d","order_by":8,"name":"Xin Jing","email":"","orcid":"","institution":"Lanzhou University","correspondingAuthor":false,"prefix":"","firstName":"Xin","middleName":"","lastName":"Jing","suffix":""},{"id":469322154,"identity":"d1352b18-9a50-43f0-ad62-7b56e67bcfb3","order_by":9,"name":"shuai ren","email":"","orcid":"","institution":"Department of Civil and Environmental Engineering, Massachusetts Institute of Technology, Cambridge, MA, USA","correspondingAuthor":false,"prefix":"","firstName":"shuai","middleName":"","lastName":"ren","suffix":""},{"id":469322155,"identity":"4afd7458-aef9-4354-942f-a2861b91fd13","order_by":10,"name":"hongbiao zi","email":"","orcid":"","institution":"State Key Laboratory of Herbage Improvement and Grassland Agro-Ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou, China","correspondingAuthor":false,"prefix":"","firstName":"hongbiao","middleName":"","lastName":"zi","suffix":""},{"id":469322156,"identity":"046e5e88-5f0d-4acf-b383-cd02a9c82fc5","order_by":11,"name":"yan shi","email":"","orcid":"","institution":"State Key Laboratory of Herbage Improvement and Grassland Agro-Ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou, China","correspondingAuthor":false,"prefix":"","firstName":"yan","middleName":"","lastName":"shi","suffix":""},{"id":469322157,"identity":"0e617fb7-1ae2-4de8-8426-2ca1cca36400","order_by":12,"name":"Tao Wang","email":"","orcid":"https://orcid.org/0000-0002-1323-8697","institution":"Institute of Tibetan Plateau Research, Chinese Academy of Sciences","correspondingAuthor":false,"prefix":"","firstName":"Tao","middleName":"","lastName":"Wang","suffix":""}],"badges":[],"createdAt":"2025-05-15 02:55:07","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6668207/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6668207/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41467-026-70187-8","type":"published","date":"2026-03-05T05:00:00+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":107131377,"identity":"14e393ad-0904-4dc3-bf2d-8c2d3a199796","added_by":"auto","created_at":"2026-04-17 07:13:04","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1052395,"visible":true,"origin":"","legend":"","description":"","filename":"main.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6668207/v1_covered_1793a514-5736-4f54-bdcf-9f881a705ab2.pdf"},{"id":84392120,"identity":"958df636-3e06-4524-9d0c-d7b9705a63fb","added_by":"auto","created_at":"2025-06-11 11:42:47","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":5070159,"visible":true,"origin":"","legend":"Supporting information for: Quantifying the trade-off between spring phenology and lethal frost risk: a global meta-analysis","description":"","filename":"appendix.docx","url":"https://assets-eu.researchsquare.com/files/rs-6668207/v1/99a1ebda20a528d9e0e72d74.docx"},{"id":84391226,"identity":"d55d7c0a-86ff-49d0-8c1e-8e727f4e21c5","added_by":"auto","created_at":"2025-06-11 11:34:47","extension":"pdf","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":89051,"visible":true,"origin":"","legend":"Reporting Summary","description":"","filename":"rs.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6668207/v1/9a21a64890e68763b5bcc320.pdf"}],"financialInterests":"There is \u003cb\u003eNO\u003c/b\u003e Competing Interest.","formattedTitle":"Quantifying the trade-off between spring phenology and lethal frost risk: a global meta-analysis","fulltext":[],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":false,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":true,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":true,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"nature-portfolio","isNatureJournal":true,"hasQc":false,"allowDirectSubmit":false,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"","title":"Nature Portfolio","twitterHandle":"","acdcEnabled":false,"dfaEnabled":false,"editorialSystem":"ejp","reportingPortfolio":"","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"","lastPublishedDoi":"10.21203/rs.3.rs-6668207/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6668207/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe timing of spring phenology reflects a critical trade-off between growing season extension and frost risk avoidance, the balance of which governs plant fitness and shapes plant species\u0026rsquo; distributions. However, few quantitative assessments of the spring lethal frost risk exist, meaning that this trade-off is largely untested empirically. Here, we present the first global meta-analysis encompassing 193 plant species across 126 study sites, finding consistently high freezing resistance (to -12\u0026deg;C) with remarkably low lethal frost risk (safety margin of 17\u0026deg;C) during spring emergence. Under projected climate warming scenarios, predicted advances in spring phenology did not affect the lethal frost risk for two plausible warming scenarios (SSP1-2.6 and SSP2-4.5), this was likely due to reduced temperature sensitivity, consistent with trade-off predictions. By contrast, in the high-warming scenario (SSP5-8.5), the freezing safety margin was predicted to expand (by ~\u0026thinsp;1\u0026deg;C), resulting in a lower lethal frost risk, this may compensate for the diminished freezing resistance predicted under warming. Our findings challenge the conventional view of simple-environmental factors moderate spring phenology and highlight the necessity of incorporating biotic factors (e.g., freezing resistance) and biotic processes (e.g., early growth versus frost risk trade-offs) into next-generation phenological models to enhance prediction accuracy under climate change.\u003c/p\u003e","manuscriptTitle":"Quantifying the trade-off between spring phenology and lethal frost risk: a global meta-analysis","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-11 11:34:42","doi":"10.21203/rs.3.rs-6668207/v1","editorialEvents":[],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"nature-communications","isNatureJournal":true,"hasQc":false,"allowDirectSubmit":false,"externalIdentity":"NCOMMS","sideBox":"Learn more about [Nature Communications](http://www.nature.com/ncomms/)","snPcode":"","submissionUrl":"https://mts-ncomms.nature.com/","title":"Nature Communications","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"ejp","reportingPortfolio":"Nature Communications","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"00f2aa8a-11a3-45da-ac93-67b6b63ebe36","owner":[],"postedDate":"June 11th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":49829221,"name":"Earth and environmental sciences/Ecology/Climate-change ecology/Phenology"},{"id":49829222,"name":"Earth and environmental sciences/Ecology/Evolutionary ecology"}],"tags":[],"updatedAt":"2026-04-17T07:12:41+00:00","versionOfRecord":{"articleIdentity":"rs-6668207","link":"https://doi.org/10.1038/s41467-026-70187-8","journal":{"identity":"nature-communications","isVorOnly":false,"title":"Nature Communications"},"publishedOn":"2026-03-05 05:00:00","publishedOnDateReadable":"March 5th, 2026"},"versionCreatedAt":"2025-06-11 11:34:42","video":"","vorDoi":"10.1038/s41467-026-70187-8","vorDoiUrl":"https://doi.org/10.1038/s41467-026-70187-8","workflowStages":[]},"version":"v1","identity":"rs-6668207","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6668207","identity":"rs-6668207","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}