Herbarium specimens reveal a cryptic invasion of tetraploid Centaurea stoebe in Europe | 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 Herbarium specimens reveal a cryptic invasion of tetraploid Centaurea stoebe in Europe Christoph Rosche, Olivier Broennimann, Andriy Novikov, Viera Mrázová, and 16 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4389565/v2 This work is licensed under a CC BY 4.0 License Status: Posted Version 2 posted You are reading this latest preprint version Show more versions Abstract Numerous plant species are expanding their native ranges due to anthropogenic environmental change. Because cytotypes of polyploid complexes show often similar morphologies, there may be unnoticed range expansions (i.e., cryptic invasions) of one cytotype into regions where only the other cytotype is native. We critically revised 13,078 herbarium specimens of diploid and tetraploid Centaurea stoebe, collected across Europe between 1790 and 2023. Based on their distribution in relictual habitats, we suggest that diploids are native across their entire European range, whereas tetraploids are native only to South-Eastern Europe and have recently expanded their range toward Central Europe. The proportion of tetraploids exponentially increased over time in their expanded but not in their native range. This cryptic invasion took predominantly place in ruderal habitats and enlarged the climatic niche of tetraploids toward a more oceanic climate. Our differentiation between native and expanded ranges conflicts with dozens of previous studies on C. stoebe . Thus, herbarium specimens can prevent erroneous assumptions on the native ranges of species, which has fundamental implications for designing research studies and assessing biodiversity trends. Moreover, we demonstrate the value of spatio-temporally explicit data in formulating and testing hypotheses regarding the superior colonization abilities of polyploids in ruderal habitats. Biological sciences/Ecology/Invasive species Biological sciences/Ecology/Biogeography Biological sciences/Plant sciences/Plant ecology Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Full Text Additional Declarations The authors declare no competing interests. Supplementary Files Roscheetal.SupportingInformation.pdf Cite Share Download PDF Status: Posted Version 2 posted You are reading this latest preprint version Show more versions 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-4389565","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":353256962,"identity":"12df465c-b267-4af2-8a3c-f572b04c586a","order_by":0,"name":"Christoph Rosche","email":"data:image/png;base64,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","orcid":"https://orcid.org/0000-0002-4257-3072","institution":"Martin Luther University","correspondingAuthor":true,"prefix":"","firstName":"Christoph","middleName":"","lastName":"Rosche","suffix":""},{"id":353256963,"identity":"fab65463-111a-4f10-982e-d72e7d0de5b2","order_by":1,"name":"Olivier Broennimann","email":"","orcid":"https://orcid.org/0000-0001-9913-3695","institution":"University of Lausanne","correspondingAuthor":false,"prefix":"","firstName":"Olivier","middleName":"","lastName":"Broennimann","suffix":""},{"id":353256964,"identity":"70a1199c-763f-4d75-95c7-10e2bd0d100e","order_by":2,"name":"Andriy Novikov","email":"","orcid":"","institution":"National Academy of Sciences of Ukraine","correspondingAuthor":false,"prefix":"","firstName":"Andriy","middleName":"","lastName":"Novikov","suffix":""},{"id":353256965,"identity":"4d13c4b6-cbe2-4278-b11d-90a021642c05","order_by":3,"name":"Viera Mrázová","email":"","orcid":"","institution":"Charles University Prague","correspondingAuthor":false,"prefix":"","firstName":"Viera","middleName":"","lastName":"Mrázová","suffix":""},{"id":353256966,"identity":"dd0b07b5-9482-45a1-b06c-61a73ee69385","order_by":4,"name":"Ganna V. 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University","correspondingAuthor":false,"prefix":"","firstName":"Marcus","middleName":"","lastName":"Lehnert","suffix":""}],"badges":[],"createdAt":"2024-05-08 13:18:00","currentVersionCode":2,"declarations":{"humanSubjects":false,"vertebrateSubjects":false,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":false},"doi":"10.21203/rs.3.rs-4389565/v2","doiUrl":"https://doi.org/10.21203/rs.3.rs-4389565/v2","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":64626341,"identity":"b599dc5d-6fbd-4fae-8da7-deac5e86bd48","added_by":"auto","created_at":"2024-09-16 18:17:42","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":735224,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eGeographic distribution of diploid (blue dots) and tetraploid \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eCentaurea stoebe \u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003eacross the European study range (n = 5,491).\u003c/strong\u003e In tetraploids, we differentiate between occurrences in the native (orange dots) and expanded ranges (red dots). In diploids, the entire study range is considered to be part of their native range. Occurrences are plotted chronologically with newer occurrences overlapping older ones. The light orange-colored area indicates the estimated native range of tetraploids. Together, the native and expanded ranges of tetraploids form our “study range” (see Methods). Note that beyond the here presented study range, diploids exhibit a substantially larger range than tetraploids. The complete distribution of both cytotypes across Eurasia is available in Supplementary Fig. 2.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-4389565/v2/f1d6c4a96b85fdff9ba45410.png"},{"id":64626595,"identity":"4f863b8d-1afe-4381-abbe-348983cb2eb2","added_by":"auto","created_at":"2024-09-16 18:25:42","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":164067,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003ePredicted proportion of tetraploid relative to all \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eCentaurea stoebe\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e records as a function of time, range, and habitat type.\u003c/strong\u003e(A) The proportion of tetraploids in the native (orange-colored) and expanded (red) ranges of tetraploids over time, irrespective of habitat type (n = 5,491). (B) The proportion of tetraploids in the expanded range of tetraploids over time, differentiated between ruderal (gray) and natural (light\u003cstrong\u003e \u003c/strong\u003egreen) habitats (n = 3,340).Model predictions are depicted using generalized additive logistic models, with solid and dashed lines denoting significant and non-significant relationships, respectively. The shaded bands show confidence intervals (1σ uncertainty). Thin gray lines indicate a proportion of tetraploids of 50%. Temporal patterns of the proportion of tetraploids in ruderal and natural habitats of the native range of tetraploids are presented in the Supplementary Fig. 5. Geographical distribution patterns of both cytotypes with respect to habitat types are presented in the Supplementary Fig. 6.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-4389565/v2/c702f4d5520dd20c3eb7b309.png"},{"id":64626344,"identity":"c89d4e57-ae5c-46bf-a361-38c6077bb9a3","added_by":"auto","created_at":"2024-09-16 18:17:42","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":69660,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTemporal dynamics of range size and niche breadth of tetraploid \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eCentaurea stoebe\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e across its expanded range, with a distinction between ruderal (gray) and natural (light green) habitats (n = 744).\u003c/strong\u003e The habitat-specific data are visualized as overlapping (dark green), not stacked, areas. (A) Range size dynamics are estimated as cumulative number of occupied 10 km × 10 km pixels over time (counting first records in distinct pixels). (B) and (C) show the temporal dynamics of the realized niches along a precipitation gradient (the first axis of the PCA, Supplementary Fig. 10) and along a temperature gradient (second PCA axis), respectively. Lines represent niche limits, with solid and dotted lines indicating significant and non-significant differences between populations from ruderal and natural habitats. Reference rectangles on the right side of the plots indicate the range size (A) and niche limits (B and C) of tetraploids in their native range (distinguished between ruderal and natural habitats and their overlap).\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-4389565/v2/f1e2115aec29db60d7d3993d.png"},{"id":64627143,"identity":"1a229303-35b5-49a0-b626-9184d84611ce","added_by":"auto","created_at":"2024-09-16 18:33:42","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":209766,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003ePredicted dispersal routes of the initial spread of tetraploid \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eCentaurea stoebe\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e across its expanded range.\u003c/strong\u003e Dispersal routes were predicted using a minimum cost arborescence algorithm (n = 951). Arrows represent these routes with line thicknesses corresponding to their bootstrap support (see legend). The inset map in the upper left corner provides context for the section of the map within Europe. The homogenously light orange-colored area represents the estimated native range of tetraploids. In the expanded ranges, climatic dissimilarity between the native and expanded ranges was assessed using a multivariate environmental similarity surface (MESS) analysis. Yellow indicates areas with climatic conditions similar to the native niche (positive MESS-values), while the beige-blue gradient illustrates the degree of dissimilarity with the climate of the native range (negative MESS-values, see legend). Mapped years denote when tetraploids were firstly reported in distinct regions: 1859: Periam, Romania; 1863: Budapest, Hungary; 1874: Devínská Nová Ves, Slovakia; 1876: Munich, Germany; 1882: Louvain, Belgium; 1908: Åhus, Sweden and Salvan, Switzerland; 1915: Oserna, Ukraine; 1925: Řepiště, Czech Republic; 1937: Paris, France; 1944: Tampere, Finland; 1981: Norderhov, Norway; 1982: Lucca, Italy; 1999: Aberford, United Kingdom.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-4389565/v2/18534b324f7b015c88c37c4a.png"},{"id":64626593,"identity":"4576298b-1e35-48eb-9be1-b7987a6a852b","added_by":"auto","created_at":"2024-09-16 18:25:42","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":119703,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003ePredictors of the spread of tetraploid \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eCentaurea stoebe\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e across its expanded range, assessed with boosted regression trees. \u003c/strong\u003e(A) Predictors of the initial spread (n = 388, cross-validation correlation (cv) = 0.49). As proxy of the initial spread, we used the residence time of tetraploids in their expanded range, i.e., the time since each of 10 km × 10 km pixels had been colonized by tetraploids. (B) Predictors of the current occurrence (n = 605, cv = 0.59). As proxy of the current occurrence, we used the predicted proportion of tetraploid relative to all \u003cem\u003eC. stoebe\u003c/em\u003e records in the timeframe between 1989 and 2023. Predictors of the two spatio-temporarily explicit response variables included three variables related to each of four categories: space (dark green), climate (black), dispersal corridors (olive), and urbanization (orange). The pie charts provide an overview of the cumulative contributions of variables categorized by data type. Bar plots show the relative importance of each variable, with upper and lower arrows indicating positive and negative relationships, respectively (\"n.s.\" denotes no significant effect). For significant predictors, plots of their linear relationships with the residence time or with the proportion of tetraploids can be found in the Supplementary Figs. 11 and 13, respectively.\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-4389565/v2/6c5a118744462d671752244b.png"},{"id":64627986,"identity":"f90b1991-6b0d-4f79-8c54-78a4138a88f3","added_by":"auto","created_at":"2024-09-16 18:49:46","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1988286,"visible":true,"origin":"","legend":"","description":"","filename":"RoscheHerbariumspecimensrevealacrypticinvasionoftetraploidCentaureastoebeinEurope.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4389565/v2_covered_c7ac6de2-9e54-49a7-b47f-8c2ed18c5e7d.pdf"},{"id":64626346,"identity":"5f32cfb4-d099-4742-b027-446edd52a291","added_by":"auto","created_at":"2024-09-16 18:17:42","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":8481214,"visible":true,"origin":"","legend":"","description":"","filename":"Roscheetal.SupportingInformation.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4389565/v2/7af36db34a71be78099c5424.pdf"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"Herbarium specimens reveal a cryptic invasion of tetraploid Centaurea stoebe in Europe","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":"","lastPublishedDoi":"10.21203/rs.3.rs-4389565/v2","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4389565/v2","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eNumerous plant species are expanding their native ranges due to anthropogenic environmental change. Because cytotypes of polyploid complexes show often similar morphologies, there may be unnoticed range expansions (i.e., cryptic invasions) of one cytotype into regions where only the other cytotype is native. We critically revised 13,078 herbarium specimens of diploid and tetraploid \u003cem\u003eCentaurea stoebe, collected across \u003c/em\u003eEurope between 1790 and 2023. Based on their distribution in relictual habitats, we suggest that diploids are native across their entire European range, whereas tetraploids are native only to South-Eastern Europe and have recently expanded their range toward Central Europe. The proportion of tetraploids exponentially increased over time in their expanded but not in their native range. This cryptic invasion took predominantly place in ruderal habitats and enlarged the climatic niche of tetraploids toward a more oceanic climate. Our differentiation between native and expanded ranges conflicts with dozens of previous studies on \u003cem\u003eC. stoebe\u003c/em\u003e.\u003cem\u003e \u003c/em\u003eThus, herbarium specimens can prevent erroneous assumptions on the native ranges of species, which has fundamental implications for designing research studies and assessing biodiversity trends. Moreover, we demonstrate the value of spatio-temporally explicit data in formulating and testing hypotheses regarding the superior colonization abilities of polyploids in ruderal habitats.\u003c/p\u003e","manuscriptTitle":"Herbarium specimens reveal a cryptic invasion of tetraploid Centaurea stoebe in Europe","msid":"","msnumber":"","nonDraftVersions":[{"code":2,"date":"2024-09-16 18:17:37","doi":"10.21203/rs.3.rs-4389565/v2","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"
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