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GUBIC: The Global Urban Biological Invasions Compendium for Plants

Li, Daijiang; Potgieter, Luke J.; Aronson, Myla F.J.; Axmanová, Irena; Baiser, Benjamin; Carboni, Marta; Celesti Grapow, Laura; Knapp, Sonja; Kühn, Ingolf; Lacerda de Matos, Ana Carolina; Vilà, Montserrat; Cadotte, Marc W.

Abstract

Urban areas are foci for the introduction of non-native plant species, and they often act as launching sites for invasions into the wider environment. Although interest in biological invasions in urban areas is growing rapidly, and the extent and complexity of problems associated with invasions in these systems have increased, data on the composition and numbers of non-native plants in urbanized areas remain scattered and idiosyncratic. We assembled data from multiple sources to create the Global Urban Biological Invasions Compendium (GUBIC) for vascular plants representing 553 urban centres from 61 countries across every continent except Antarctica. The GUBIC repository includes 8140 non-native plant species from 253 families. The number of urban centres in which these non-native species occurred had a log-normal distribution, with 65.2% of non-native species occurring in fewer than 10 urban centres. Practical implications: The dataset has wider applications for urban ecology, invasion biology, macroecology, conservation, urban planning and sustainability. We hope this dataset will stimulate future research in invasion ecology related to the diversity and distributional patterns of non-native flora across urban centres worldwide. Further, this information should aid the early detection and risk assessment of potential invasive species, inform policy development and assist in setting management priorities.

Full text

Ecol Solut Evid. 2025;6:e70020.   | 1 of 11 https://doi.org/10.1002/2688-8319.70020 wileyonlinelibrary.com/journal/eso3 Received:12December2024 | Accepted:25January2025 DOI: 10.1002/2688-8319.70020 DATA ARTICLE GUBIC: The global urban biological invasions compendium for plants Daijiang Li1 | Luke J. Potgieter2,3 | Myla F. J. Aronson4 | Irena Axmanová5 | Benjamin Baiser6 | Marta Carboni7 | Laura CelestiGrapow8 | Sonja Knapp9,10 | Ingolf Kühn9,10,11 | Ana Carolina Lacerda de Matos12 | Zdeňka Lososová5 | Flavia A. MontañoCentellas13 | Petr Pyšek14,15 | David M. Richardson3 | Lauren B. Trotta6 | Rafael D. Zenni12 | Sarel S. Cilliers16 | Bruce D. Clarkson17 | Amy J. S. Davis18 | Rebecca W. Dolan19 | Marcin K. Dyderski20 | Franz Essl21 | Orou G. Gaoue22 | Joanne Gui2 | Charly Géron23,24 | Gustavo Heringer12,25,26 | Cang Hui27,28 | Anzar A. Khuroo29 | Stefan Klotz9 | Peter M. Kotanen30 | Holger Kreft31,32 | Frank A. La Sorte33,34 | Jonas J. Lembrechts35 | Bernd Lenzner21 | Christopher A. Lepczyk36 | Scott MacIvor2 | Cristina MartínezGarza37 | Akira S. Mori38 | Charles Nilon39 | Jan Pergl14 | Stefan J. Siebert16 | Alyona S. Tretyakova40 | Toby P. N. Tsang2 | Kei Uchida41 | Mark van Kleunen18,42 | Montserrat Vilà43,44 | HuaFeng Wang45 | Patrick Weigelt46 | Peter Werner47 | Nicholas S. G. Williams48 | Marten Winter10 | Marc W. Cadotte2 This is an open access article under the terms of the CreativeCommonsAttribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. ©2025TheAuthor(s).Ecological Solutions and EvidencepublishedbyJohnWiley&SonsLtdonbehalfofBritishEcologicalSociety. DaijiangLiandLukeJ.Potgietercontributedequallytothiswork. For affiliations refer to page 9. Correspondence Daijiang Li Email:[email protected] LukeJ.Potgieter Email:[email protected] MarcW.Cadotte Email:[email protected] Funding information NaturalSciencesandEngineering Research Council of Canada, Grant/AwardNumber:#RGPIN- 202203579 and 386151; Deutsche Forschungsgemeinschaft, Grant/ AwardNumber:264740629,FZT118 and202548816;NationalScience Abstract 1. Urban areas are foci for the introduction of nonnative plant species, and they oftenactaslaunchingsitesforinvasionsintothewiderenvironment.Although interest in biological invasions in urban areas is growing rapidly, and the extent and complexity of problems associated with invasions in these systems have increased, data on the composition and numbers of nonnative plants in urbanized areas remain scattered and idiosyncratic. 2. WeassembleddatafrommultiplesourcestocreatetheGlobalUrbanBiological InvasionsCompendium(GUBIC)forvascularplantsrepresenting553urbancentresfrom61countriesacrosseverycontinentexceptAntarctica. 3. TheGUBICrepositoryincludes8140non-nativeplantspeciesfrom253families. The number of urban centres in which these nonnative species occurred had a 2 of 11 | LI et al. 1 | INTRODUCTION Urban areas, characterized by their high human population densityandextensivelandscapemodification,presentuniqueopportunities for the establishment and spread of nonnative species. The convergence of global trade, transportation networks, modified microclimates and humanmediated disturbances in urban areas facilitates the introduction and proliferation of nonnative species(Gallardoetal.,2016;Potgieteretal.,2024).Urbanplant invasions can have profound ecological, economic and social impacts due to altered ecosystem services, impacts on human health and costs incurred from management efforts (Heringer et al., 2024;Potgieteretal.,2017).However,thereisalackof foundational data on which species occur in urban centres globally. This data gap limits our ability to assess the potential threats nonnative plants pose to urban ecosystems and the services they mightprovide(Milanovićetal.,2020),withcurrentknowledgeremaining geographically heterogeneous and focused on only a few well-studiedtaxa(Vazetal.,2018). Frameworks for understanding and managing urban plant invasionsarelessfrequentlystudiedthaninotherhabitats(but see Gaertner et al., 2016;Potgieter&Cadotte,2020).Whileexisting frameworks integrate concepts from landscape ecology, population biology and socioecological systems, they are limited in number and scope, highlighting the need for further development to facilitate a better understanding of the mechanisms that drive invasions in urban areas as well as options for managing them.Managersinurbanareasfaceuniquechallengesduetothe interplay between the built environment and complex socioeconomic factors, which can significantly alter ecosystem conditions. However,thesechallengeshaveonlyrecentlybeenincorporated into models to predict urban invasion dynamics and impacts and identifyappropriatemanagementstrategies(Gaertneretal.,2016; Potgieteretal.,2022). Despite these advances, empirical studies on urban biological invasions remain limited, particularly in terms of taxonomic coverageandspatialscale(Cadotteetal.,2017).Mostempirical studies have focused on the ecology of particular nonnative species within small urban areas. This narrow focus limits the generalizability of findings across different organisms and urban contexts. Althoughnumerousregionalandcity-specificinventoriesofnon- native species exist, these are often from uncoordinated efforts carried out independently by research groups focusing on particularresearchquestions.Therefore,thesediverseresourceslack harmonization of collection methods, taxonomy and sampling effort, making them challenging to be easily used. Moreover, because some of this work is developed in collaboration with city practitioners and managers, many studies are published in the greyliteratureandonlyavailableinnon-Englishlanguages,limiting theiraccessibility.Whilethesebiologicalinventoriesarecrucial to advancing our understanding of urban biological invasions at the city and regional levels, a comprehensive global dataset documenting the nonnative flora in urban areas around the world isrequiredtounderstandtheroleofurbanareasinshapingthe patternsofplant invasionsandtheunderlying processes.Here, we unify this diverse body of knowledge and present a global repository of nonnative flora in urban centres around the globe. This repository serves as a valuable resource for improving our understanding of urban nonnative floras by providing essential data, fostering collaboration, informing management and policy and facilitating coordinated global responses to the challenges they present. 2 | METHODS AND MATERIALS Tocompilealistofnon-nativeplantspeciesinurbanareas(see Section2.1.2 for the methods used to delineate urban boundaries)globally,wecombinedmultipledatasources.Thisapproach allows for the application of standardized selection and inclusion criteria over multiple individual datasets, resulting in a harmonized and consistent dataset across urban areas and regions. lognormal distribution, with 65.2% of nonnative species occurring in fewer than 10 urban centres. 4. Practical implications: The dataset has wider applications for urban ecology, invasionbiology,macroecology,conservation,urbanplanningandsustainability.We hope this dataset will stimulate future research in invasion ecology related to the diversity and distributional patterns of nonnative flora across urban centres worldwide. Further, this information should aid the early detection and risk assessment of potential invasive species, inform policy development and assist in setting management priorities. KEYWORDS Alienspecies,biodiversitychange,biologicalinvasions,cities,naturalizedspecies,non-native plants, urbanization Foundation,Grant/AwardNumber: DEB-2213567;TheItalianMinistryof UniversityandResearch,Grant/Award Number:CN00000033;CNPq-Brazil, Grant/AwardNumber:302643/2022-2; CzechScienceFoundation,Grant/Award Number:19-28491Xand25-15190S; CzechAcademyofSciences,Grant/Award Number:RVO67985939 Handling Editor:HollyJones 26888319, 2025, 1, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1002/2688-8319.70020 by Readcube (Labtiva Inc.), Wiley Online Library on [02/05/2025]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License | 3 of 11 LI et al. Weincludedonlyestablishednon-nativeplantspecies,whichare those with selfsustaining populations, also commonly referred to asnaturalized(Blackburnetal.,2011; Richardson et al., 2000; see Section2.1.5). 2.1 | Data acquisition and compilation 2.1.1 | Datasource1:GlobalUrbanBiological Invasions Consortium An international workshop to address biological invasions in urbanecosystemswashostedbytheCentreforInvasionBiology in Stellenbosch, South Africa, in November 2016 (Gaertner et al., 2017).ThisworkshopledtothecreationoftheGlobal Urban BiologicalInvasionsConsortium, which hosted a coordinating meetinginJune2019thatbroughttogethermorethan70researchers from14countriesfromallcontinentsexceptAntarctica.Oneofthe prioritized activities was to compile lists of nonnative plant speciesforurbanareas.Aworkinggroup“SynthesizingGlobalUrban BiologicalInvasionKnowledge”(sGUBIK,fundedbysDiv,thesynthesiscentreofiDiv,theGermanCentreforIntegrativeBiodiversity Research) was later established in September 2023 to synthesize these global data and examine the patterns and mechanisms driving nonnative plant species' invasions in urban areas. Wecompileddatausingthefollowing approaches.First,we sentarequesttoover150membersoftheGlobalUrbanBiological Invasions Consortium in 2019 to upload datasets for any urban taxatoaSharePointrepositoryattheUniversityofToronto.The cut-offforthedatarequestwasDecember2021.Second,during AugusttoNovember2019,wesearchedthepublishedliteraturein English,PortugueseandSpanishaswellastheDryaddatarepository(www. datad ryad. org)forstudiesanddatasetscontainingspecies lists for urban areas around the world, using keywords such as ‘alien’, ‘animal’, ‘builtup’, ‘city’, ‘urban*’, ‘nonnative’, ‘exotic’ and ‘plant’. These approaches yielded urban datasets that encompassed various taxa and spatial scales, incorporating demographic, environmental and taxon-specific information. Additionally, we includedtheUrbanBiodiversityResearchCoordinationNetwork (UrBioNet) dataset, a large multi-city compilation (Aronson et al., 2014),featuring14,240spontaneousplantspecies(i.e.not cultivatedorplanted),ofwhich4241areidentifiedasnon-native, derived from published surveys across 110 urban areas in five biogeographic regions. To ensure consistency across the datasets, we standardized city and country names by resolving variations in spelling and correcting potential typographical errors. In instances where multiple urbancentreswithinthesamecountrysharedthesamename(e.g. Madison,Wisconsinvs.Madison,IndianaintheUnitedStates),we excluded these entries from the database if it was not possible to unambiguously determine the specific city to which the data pertained. Given that most data lacked spatially explicit coordinates, precise delineations of city boundaries were unavailable. Asaresult,datasetscollectedfromdatacontributors,repositories or the literature were generally treated as representing areas surrounding the urban centres rather than being confined to specific urban boundaries. 2.1.2 | Datasource2:GlobalBiodiversity Information Facility BeforeextractingoccurrencedataforeachurbanareafromtheGlobal Biodiversity Information Facility (GBIF), we delineated the boundariesofurbanareas.Weusedtheglobalurbancentresdataprovided bytheGlobalHumanSettlementLayer(GHSL,Pesaresietal.,2019, https:// ghsl. jrc. ec. europa. eu/ ucdb2 018Ov erview. php), which definesurbancentresascontiguous1 km2 grid cells with a population density of at least 1500 inhabitants per km2ofpermanentland(areas that are consistently above water and exclude bodies of water, such asoceans,seas,largeriversandlakes)orwithmorethan50%built-up surface shared on permanent land and with at least 50,000 inhabitantsintheclusterwithsmoothedboundariesandsmallgaps(<15 km2) filled. Overall, there are 13,189 unique urban centres worldwide. Subsequently,smaller,nearbyurbancentreslocatedwithina5 kmradius of the larger urban centres were integrated into the larger one, as these proximally situated centres are close enough to be considered a single urban entity and often are considered part of the metropolitan area.Werefrainedfromfurthermergingsmallercentresthat,although withina5 kmradiusofthepreviouslymergedsmallercentres,were situatedbeyondthe5 kmboundaryfromthelargerurbancentre.This processresultedin11,621uniqueurbancentresglobally. InAugust2023,wequeriedGBIFanddownloadedplantoccurrence records from each urban centre to compile the flora of these urbanareas(seeTable S1fortheDOIsofdownloadeddatasets).The initial download comprised over 500 million records. We cleaned the GBIF data of each of the urban centres by removing records with common issues such as erroneous coordinates using R package ‘CoordinateCleaner’(Zizkaetal.,2019).Wealsoremovedallrecords with identification above species level, fossil specimens, preserved specimens, living specimens and those with locality uncertainty greaterthan30 kmorwithina500 mvicinityofbiodiversityinstitutions,botanicgardens,zoos,museums,GBIFheadquarters,etc. 2.1.3 | Qualitycontrolandmergingofdata Beforemergingdatafromsources1and2,weconductedpreliminary filteringof thesedatasets.ForeachurbancentrewithGBIF data, we used the number of observations of each species as a proxy fortheabundanceofthatspecies.Wecalculatedobservedspecies richnessandestimatedspeciesrichnessusingtheChao1equation, whichincorporatessingletonsanddoubletons(i.e.speciesobserved onlyonceortwice): (1) Estimated richness = Observed richness + S 2∕( 2D ) 26888319, 2025, 1, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1002/2688-8319.70020 by Readcube (Labtiva Inc.), Wiley Online Library on [02/05/2025]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License 4 of 11 | LI et al. where S represents the number of singletons and D is the number of doubletons (Hsieh & Chao, 2016). We also determined the sample coverage percentage, a measure of sample completeness, based on the rarefied estimate of the total number of individuals in each urban centreusingtheRpackage‘iNEXT’forrarefaction(Chaoetal.,2014; Hsiehetal.,2024). WeconsideredanurbancentretohaverobustGBIFdataif:(1)it hadover1000observedplantspecies;(2)thecommunitysamplecoverage was >90%;and(3)theobservedspeciesrichnesswasgreater than75%oftheestimatedspeciesrichness.Weusedthesecriteria tobalancethenumberofretainedurbancentresanddataquality.For data source 1, if an urban centre had more than 300 plant species, we retaineditandfurtherintegrateditwithdatasource2(GBIFdata)of thaturbancentreregardlessoftheGBIFdataquality.Ifanurbancentre had fewer than 300 species from data source 1 and did not have adequateGBIFdatacoverage,weremovedthaturbancentrefromour database. If an urban centre had fewer than 300 species from data source 1but hadadequateGBIF data coverage (i.e. met the above threecriteria),weretainedbothdatasourcesforthaturbancentre.We removedthoseurbancentreswithonlyGBIFdatathatdidnotmeet thethreecriteriaabove(seeFigure 1foraschematicworkflow).Like thecriteriaweusedfortheGBIFdata,weselected300specieshere tobalancethenumberofurbancentresandtheirdataqualityafter carefully explored our datasets. The final database included 553 urban centres(Figure 2).Foreachoftheseurbancentres,wederivedalistof established nonnative plant species using the merged data sources. 2.1.4 | Standardizespeciesnames We standardized species and family names against the World Checklist of Vascular Plants (WCVP, Govaerts, 2024) for the mergeddatabaseusingtheRpackagerWCVP(version1.0.3,Brown et al., 2023).WeselectedWCVPasitrepresentsoneofthemostcomprehensive and up-to-date taxonomic resources available (Grenié et al., 2022).WCVPalsoservesasthetaxonomicbackboneforthe mostrecentversionoftheGlobalNaturalizedAlienFlora(GloNAF), whichwasupdatedfollowingvanKleunenetal.(2019).GloNAFwas used to determine whether a species is nonnative in a particular regionwhereanurbancentrewaslocated(seeSection2.1.5below). Notethatspecieswith“unplacednames”(n = 65acrossallspecies) orhasnotmatchfromWCVPwereexcludedfromthefinaldataset (h t t p s : // p o w o . s c i e n c e . k e w . o r g / a b o u t - w c v p # u n p l a c e d n a m e s ), reflecting the challenges in our current taxonomic knowledge of plants worldwide.Wealsomergedsubspeciesorvarietiestothemainspecies and only kept binomial species names in the final database. 2.1.5 | Cross-validationtodeterminethestatusof species Todistinguishbetweenestablished(naturalized)andnativeorcasual species(i.e.thosethatmightflourishandevenreproduceoccasionally in an area but which do not form selfreplacing populations; FIGURE 1 SchematicfigureshowingtheworkflowofthecompilationoftheGlobalUrbanBiologicalInvasionsCompendiumdatabase. UrBioNet:TheUrbanBiodiversityResearchCoordinationNetwork.GHSL,TheGlobalHumanSettlementLayer;GBIF,TheGlobal BiodiversityInformationFacility;WCVP,TheWorldChecklistofVascularPlants;GloNAF,GlobalNaturalizedAlienFlora. Literature search Contributed data (e.g., UrBioNet) Data repository (e.g., Dryad) Data source 1 If an urban centre has < 300 species and no good GBIF data GHSL global urban centers Data source 2 Query GBIF Data integration Data cleanup Drop this urban centre < 300 species and good GBIF data or has > 300 species If an urban centre has Keep this urban centre and integrated with data source 2 Keep this urban centre and integrated with data source 1 if available Good GBIF data or does not have good GBIF data but has good data from source 1 No good GBIF data and has no good data from source 1 Drop this urban centre Integrated flora list of urban centres Standardize species names with WCVP Determine status of species within each urban centre using GloNAF GUBIC data: list of non-native species of 553 urban centers / cities across the world 26888319, 2025, 1, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1002/2688-8319.70020 by Readcube (Labtiva Inc.), Wiley Online Library on [02/05/2025]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License | 5 of 11 LI et al. FIGURE 2 Geographicdistributionofurbancentresacrosstheworld(panela;n = 553)andEurope(panelb)andthenumberof establishednon-nativeplantspeciestheycontain(colouredpoints). # of non−native species (log10) 1.52.0 2.53.0 (a) (b) FIGURE 3 Thedistributionoffamilysizesforthe253establishednon-nativeplantfamiliesinthedataset.Themainplotcontainsthetop 20 families which together account for 61.6% of all established nonnative plant species in our dataset. The numbers after the family names represent the approximate number of total accepted species of each family. The embedded plot presents the distribution of the number of nonnative plant species across all families. 853 784 645 359 248 227 213 167 157 148 143 135 135 134 128 125 117 105 96 96 Convolvulaceae (2496) Myrtaceae (7171) Asparagaceae (4024) Polygonaceae (2143) Apiaceae (4998) Boraginaceae (4332) Iridaceae (3007) Euphorbiaceae (7591) Malvaceae (6452) Plantaginaceae (2985) Caryophyllaceae (4645) Solanaceae (3157) Amaranthaceae (3021) Cyperaceae (7094) Brassicaceae (5382) Lamiaceae (10042) Rosaceae (6851) Fabaceae (28400) Poaceae (13919) Asteraceae (45021) 0 200 400 600 800 Number of non−native species 0 200 400 600 800 Number of non−native species 26888319, 2025, 1, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1002/2688-8319.70020 by Readcube (Labtiva Inc.), Wiley Online Library on [02/05/2025]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License 6 of 11 | LI et al. Richardson et al., 2000)locatedinaspecificurbancentre,weused theGloNAFdatabaseasitprovidesthemostupdatedinformation of naturalized plant species across the world. For each urban centre, weusedthedelineatedboundariesprovidedbytheGHSL.Foreach species listed within an urban centre, we classified the species as nonnative to that urban centre if its polygon intersected with the species'naturalizedorinvasiverange.Wealsocross-referencedall species with local checklists of nonnative plant species validated by experts(Kalusováetal.,2024).Therefore,forthoseurbancentres (mostlyinEurope),thelistsofnaturalizedspecieswereslightlydifferentfromthosebasedonGloNAFalone. 3 | GENERAL PATTERNS We present a global urban non-native flora for 553 cities from 61 countries across every continent with permanent human settlements (Figure 2). These data are, however, biased towards EuropeanandNorthAmericanurbancentres,whichtogetheraccount for 80.8% of all nonnative species, and 82.2% of all records within our database across the world, respectively. Our global repository includes 8140 established nonnative plant species from 253families(Figure 3).Mostfamiliescontainfewspecies,with73 familieseachcontaining20ormorenon-nativespecies(Figure 3). Asteraceae,Poaceae,FabaceaeandRosaceaecontainaboutone- thirdofallspecies(n = 2641;Table 1).Themostwidespreadnon- native plant species can be found in Table 1; the top 20 urban centres and countries with the greatest number of nonnative plant species in our database can be found in Table 2.Ararefactionof speciesoccurrencesacrossurbancentres(Figure 4)showsthatwe are approaching an asymptote with our sample of 553 urban centres.However,thesamplingcurvealsosuggeststhatmoreurban floral sampling is needed, especially from regions with sparse data (e.g.SouthAsia,northernSouthAmerica). TABLE 1 Themostwidespread(top30)establishednon-nativeplantspeciesinurbancentres(n = 553)acrosstheworld.Notethatthislist wasderivedfromdifferentsamplingeffortsandhasabiasinfavourofnon-nativespeciesinEuropeanandNorthAmericanurbancentres. Scientific name Family Number of urban centres Number of countries Number of GBIF records Erigeron canadensis Asteraceae 469 47 64,760 Veronica persica Plantaginaceae 451 41 41,176 Oxalis corniculata Oxalidaceae 434 48 23,721 Datura stramonium Solanaceae 410 46 12,531 Robinia pseudoacacia Fabaceae 404 41 44,657 Syringa vulgaris Oleaceae 393 29 21,767 Amaranthus retroflexus Amaranthaceae 381 41 8602 Galinsoga quadriradiata Asteraceae 376 40 18,509 Ailanthus altissima Simaroubaceae 369 35 50,732 Medicago sativa Fabaceae 369 39 24,969 Prunus cerasifera Rosaceae 369 26 26,924 Aesculus hippocastanum Sapindaceae 367 24 35,349 Reynoutria japonica Polygonaceae 367 29 88,542 Cymbalaria muralis Plantaginaceae 366 33 31,151 Matricaria discoidea Asteraceae 363 33 26,254 Melissa officinalis Lamiaceae 359 25 20,768 Buddleja davidii Scrophulariaceae 356 33 45,122 Lunaria annua Brassicaceae 340 19 12,135 Galinsoga parviflora Asteraceae 337 43 7306 Rosa rugosa Rosaceae 334 22 18,499 Vinca major Apocynaceae 326 23 11,597 Helianthus tuberosus Asteraceae 320 34 4244 Tanacetum parthenium Asteraceae 320 27 9527 Lepidium draba Brassicaceae 315 29 14,831 Acer negundo Sapindaceae 313 36 17,459 Lysimachia punctata Primulaceae 312 15 10,279 Brassica napus Brassicaceae 307 25 5376 Impatiens glandulifera Balsaminaceae 301 25 35,332 Lepidium didymum Brassicaceae 301 31 13,459 Prunus laurocerasus Rosaceae 298 22 48,188 26888319, 2025, 1, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1002/2688-8319.70020 by Readcube (Labtiva Inc.), Wiley Online Library on [02/05/2025]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License | 7 of 11 LI et al. 4 | USAGE NOTES In forming a dataset of this magnitude, we made several simplifying decisionsandrecognizethatlimitationsareinevitable.Someissues to be cognizant of for analysis and interpretation include: 1. Our definition of urbanized areas delineated contiguous areas. Becauseofthisdefinition,someurbanizedareasspanmultiple regions or municipalities and form contiguous land areas. In these cases, the urbanized region is referred to as the largest administrative centre; for example, Guangzhou, China includes TABLE 2 Thetop20urbancentres(left)andthetop20countries(right)withthegreatestnumberofnon-nativeplantspecies.Notethat theselistsareskewedtowardsEuropeanandNorthAmericanurbancentres(seeFigure 2).Thenumberspresentedforsomecountries (e.g.France)alsoincludednon-nativeplantspeciesfromtheiroverseasurbancentres. Urban centre Number of established nonnative species Country Number of established nonnative species New York 1663 UnitedStatesofAmerica 4409 LosAngeles 1534 Australia 2596 Sydney 1486 France 2187 Philadelphia 1455 NewZealand 1561 Melbourne 1450 Canada 1476 WashingtonD.C. 1414 Russia 1251 Auckland 1310 Japan 1154 Boston 1300 Mexico 1142 SanJose(USA) 1231 Germany 1123 Tijuana 1066 UnitedKingdom 986 St.Louis 1058 Switzerland 966 Tokyo 1038 SouthAfrica 947 London(UK) 1014 Spain 916 Christchurch 1009 Belgium 906 Adelaide 995 Netherlands 860 Brisbane 994 Sweden 832 Portland(OR,USA) 990 Denmark 800 Moscow 924 Norway 792 Chicago 895 Portugal 596 Perth 858 Brazil 588 FIGURE 4 Rarefactioncurveofthe number of nonnative plant species in 553 urban centres. 0 2500 5000 7500 0 200 400600 800 Number of urban centres Number of non−native species Rarefaction Extrapolation 26888319, 2025, 1, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1002/2688-8319.70020 by Readcube (Labtiva Inc.), Wiley Online Library on [02/05/2025]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License 8 of 11 | LI et al. Foshan. In some cases, contiguous urbanized areas span larger administrative areas and even countries. For example, Detroit, Michigan,USA,notonlyincludesneighbouringcitiesinMichigan, like Dearborn, but also the Canadian city of Windsor. 2. Whilemostrecordedspeciesinourdatasetcanbeconfirmedas established, the status of some species could not be definitively verified with our methodology. Additionally, the dataset might include nonestablished nonnative plant species or intentionally cultivated individuals that were not fully distinguishable from naturallyoccurringrecords.Asaresult,thedatashouldbeinterpreted cautiously, particularly when comparing nonnative species richness at broader spatial scales, such as across countries, ratherthanatthecitylevel. Speciesin our dataset withwidespread occurrences across multiple urban centres are likely to be established, whereas species recorded in only one urban centre mightrequirefurtherscrutiny.Werecommendthatusersconsider including these singleton records in sensitivity analyses to assess the robustness of their results. Therefore, the numbers of non-nativespeciesreportedhere(e.g.Table 2)areinsomecases higher than those reported for individual countries in recent studies(Kalusováetal.,2024;Pyšeketal.,2017). 3. The combination of these many individual datasets means that our list is subject to numerous methodological differences, from lists being built from herbarium specimens to those observed duringdirectsampling.Becauseourgoalistocompileanon-native flora of urban centres, these limitations do not significantly affect our dataset. 4. ThedataextractedfromGBIFincludegeographicallybiasedand incomplete sampling, and species counts derived from these data should not be considered exhaustive despite our strict criteria listed above. For example, many urban centres in China included fewer than 100 nonnative species in our database (Figures 2 and 5a),whicharelikelyunderestimates.Analyses of richness and diversity should include rarefaction or some otherwayofaccountingforunequalsamplingasthenumberof nonnative species increased with the number of observations (Figure 5b).Notably,manyurbancentresfromtheGlobalSouth (e.g.India;Figure 2)wereabsentfromourdatabaseduetothe paucity of available data. 5 | CONCLUSION Thedatabasepresentedhererepresentsauniqueandvaluableresource for addressing a wide range of basic and applied ecological questions,particularlythoserelatedtobiologicalinvasions.Thisresource can support hypothesis testing at the macroand global scale (e.g.bioticresistanceorinvasiondebt).Itcanalsobeusedtomodel nonnative plant species invasions, underscoring its utility not only in scientific research but also in conservation planning and practice. Lastly, it has the potential to guide more informed decisionmaking in biodiversity conservation, ecosystem restoration, environmental sustainability and invasive species management across diverse ecological, biogeographical and urban contexts. AUTHOR CONTRIBUTIONS The Global Urban Biological Invasions Consortium, led by Marc Cadotte and including all coauthors, conceived the initial idea, which wasfurtherdevelopedbythesGUBIKworkinggroup(DaijiangLi,Luke J. Potgieter, Myla F. J. Aronson, Irena Axmanová, Benjamin Baiser, MartaCarboni,LauraCelesti-Grapow,SonjaKnapp,IngolfKühn,Ana Carolina Lacerda de Matos, Zdeňka Lososová, Flavia A. Montaño- Centellas,PetrPyšek,DavidM.Richardson,LaurenB.Trotta,RafaelD. ZenniandMarcW.Cadotte).LukeJ.Potgieterledthedatacollection FIGURE 5 (a)Distributionofthenumberofnon-nativeplantspeciesinurbancentres(n = 553)across61countriesand(b)therelationship between the number of nonnative plant species and the number of nonnative species occurrence records in that dataset. 0 25 50 75 0 500 1000 1500 Number of non−native plant species Number of urban centres (a) 30 100 300 1000 101102103104105106 Number of non−native species observations Number of non−native species (b) 26888319, 2025, 1, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1002/2688-8319.70020 by Readcube (Labtiva Inc.), Wiley Online Library on [02/05/2025]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License | 9 of 11 LI et al. effort, gathering information from the literature and contributors, who were invited to join as coauthors if they provided additional contributions to the manuscript. Daijiang Li led the data compilation from GBIF.ThesGUBIKteamsupporteddataintegrationandstandardization.LukeJ.PotgieterandDaijiangLidraftedthemanuscript,withall coauthors contributing to its editing and revision. AFFILIATIONS 1DepartmentofEcologyandEvolutionaryBiology,UniversityofArizona, Tucson,Arizona,USA;2DepartmentofBiologicalSciences,Universityof Toronto-Scarborough,Toronto,Ontario,Canada;3Centre for Invasion Biology,DepartmentofBotany&Zoology,StellenboschUniversity, Stellenbosch,SouthAfrica;4DepartmentofEcology,Evolutionand NaturalResources,Rutgers,TheStateUniversityofNewJersey,New Brunswick,NewJersey,USA;5DepartmentofBotanyandZoology,Faculty ofScience,MasarykUniversity,Brno,CzechRepublic;6Department of WildlifeEcologyandConservation,UniversityofFlorida,Gainesville, Florida,USA;7DepartmentofSciences,RomaTreUniversity,Rome,Italy; 8DepartmentofEnvironmentalBiology,SapienzaUniversityofRome, Rome, Italy; 9DepartmentofCommunityEcology,HelmholtzCentrefor EnvironmentalResearch-UFZ,Halle(Saale),Germany;10German Centre forIntegrativeBiodiversityResearch(iDiv),Halle-Jena-Leipzig,Leipzig, Germany; 11GeobotanyandBotanicalGarden,MartinLutherUniversity Halle-Wittenberg,Halle(Saale),Germany;12DepartamentodeEcologiae Conservação, Instituto de Ciências Naturais, Federal University of Lavras, Lavras,MinasGerais,Brazil;13DepartmentofBiologicalSciences,Louisiana StateUniversity,BatonRouge,Louisiana,USA;14Department of Invasion Ecology,InstituteofBotany,CzechAcademyofSciences,Průhonice, Czech Republic; 15DepartmentofEcology,FacultyofScience,Charles University,Prague,CzechRepublic;16UnitforEnvironmentalSciences andManagement,North-WestUniversity,Potchefstroom,SouthAfrica; 17EnvironmentalResearchInstitute,UniversityofWaikato,Hamilton,New Zealand;18EcologyGroup,DepartmentofBiology,UniversityofKonstanz, Konstanz,Germany;19FriesnerHerbarium,ButlerUniversity,Indianapolis, Indiana,USA;20InstituteofDendrology,PolishAcademyofSciences, Kórnik,Poland;21DivisionofBioInvasions,GlobalChange&Macroecology, DepartmentofBotanyandBiodiversityResearch,UniversityofVienna, Vienna,Austria;22DepartmentofEcologyandEvolutionaryBiology, UniversityofTennessee,Knoxville,Tennessee,USA;23Biodiversityand Landscape,TERRAResearchCentre,GemblouxAgro-BioTech,Universityof Liège,Gembloux,Belgium;24PlantsandEcosystems,UniversityofAntwerp, Wilrijk,Belgium;25Nürtingen-GeislingenUniversity(HfWU),Nürtingen, Germany; 26ThünenInstitute,InstituteofBiodiversity,Braunschweig, Germany; 27CentreforInvasionBiology,DepartmentofMathematical Sciences,StellenboschUniversity,Stellenbosch,SouthAfrica;28African InstituteforMathematicalSciences,NationalInstituteforNational InstituteforTheoreticalandComputationalSciences,CapeTown,South Africa;29CentreforBiodiversity&Taxonomy,DepartmentofBotany, UniversityofKashmir,Srinagar,JammuandKashmir,India;30Department ofEcologyandEvolutionaryBiology,UniversityofTorontoMississauga, Mississauga, Ontario, Canada; 31Biodiversity,Macroecology&Biogeography, University of Göttingen, Göttingen, Germany; 32CentreofBiodiversityand SustainableLandUse(CBL),UniversityofGöttingen,Göttingen,Germany; 33DepartmentofEcologyandEvolutionaryBiology,YaleUniversity,New Haven,Connecticut,USA;34CenterforBiodiversityandGlobalChange, YaleUniversity,NewHaven,Connecticut,USA;35Ecology&Biodiversity, DepartmentofBiology,UtrechtUniversity,Utrecht,TheNetherlands; 36CollegeofForestry,WildlifeandEnvironment,AuburnUniversity,Auburn, Alabama,USA;37CentrodeInvestigaciónenBiodiversidadyConservación, UniversidadAutónomadelEstadodeMorelos,Cuernavaca,Mexico; 38ResearchCenterforAdvancedScienceandTechnology,TheUniversityof Tokyo,Tokyo,Japan;39SchoolofNaturalResources,UniversityofMissouri, Columbia,Missouri,USA;40Ekaterinburg,Russia;41FacultyofEnvironmental Studies,TokyoCityUniversity,Tokyo,Japan;42ZhejiangProvincialKey LaboratoryofPlantEvolutionaryEcologyandConservation,Taizhou University, Taizhou, China; 43DepartamentodeBiologíaVegetalyEcología, UniversidaddeSevilla,Sevilla,Spain;44EstaciónBiológicadeDoñana (EBD-CSIC),Sevilla,Spain;45KeyLaboratoryofTropicalBiologicalResources ofMinistryofEducation,SchoolofLifeandPharmaceuticalSciences,Hainan University,Haikou,China;46DepartmentofEnvironmentalScience,Radboud InstituteforBiologicalandEnvironmentalSciences,RadboudUniversity, Nijmegen, The Netherlands; 47InstituteforHousingandEnvironment, Darmstadt, Germany and 48SchoolofAgricultureFoodandEcosystem Science,UniversityofMelbourne,Parkville,Victoria,Australia FUNDING INFORMATION This paper is a joint effort of the working group sGUBIK kindly supportedbysDiv,theSynthesisCentreoftheGermanCentrefor IntegrativeBiodiversityResearch(iDiv)Halle-Jena-Leipzig,funded by the German Research Foundation (FZT 118, 202548816). The Global Urban Biological Invasion Consortium was initially funded bytheConnaughtGlobalChallengesAward,theOfficeoftheVice- PresidentInternational,theSchoolofGraduateStudies,University of Toronto, and the Office of the Vice-Principal Research at the UniversityofTorontoScarborough.L.J.P.andD.M.R.acknowledge support from the Centre for Invasion Biology and Stellenbosch University.D.L.wassupportedby USNSFDEB-2213567.M.W.C. wassupportedbytheNaturalSciencesandEngineeringResearch CouncilofCanada(#386151).Z.L.andP.P.weresupportedbygrant no.25-15190S(CzechScienceFoundation)andlong-termresearch developmentprojectRVO67985939(CzechAcademyofSciences). Z.L.andI.A.weresupportedbyEXPROgrantno.19-28491X(Czech Science Foundation). M.C. acknowledges the support of NBFC, fundedbytheItalianMinistryofUniversityandResearch,PNRR, Missione4Componente2,“Dallaricercaall'impresa”,Investimento 1.4, Project CN00000033. R.D.Z. acknowledges the support of CNPq-Brazil (302643/2022-2). M.v.K., A.D., and M.W. acknowledge funding of the German Research Foundation (M.v.K., A.D.: 264740629,M.W.via iDiv). P.M.K.was supportedby the Natural Sciences andEngineering ResearchCouncil ofCanada(Discovery Grant #RGPIN-2022-03579). M.V. by (PID2021-122690OB-I00) funded by MCIN/AEI/10.13039/501100011033/FEDER, UE. G.H. acknowledgesthesupportofAlexandervonHumboldtFoundation andCoordenaçãodeAperfeiçoamentodePessoaldeNívelSuperior– Brasil(Capes)–Financecode001.M.K.D.acknowledgesthesupport oftheInstituteofDendrology,PolishAcademyofSciences.A.S.M wassupportedbytheMitsui&Co.,Ltd.EnvironmentFund(funding no.R17-0062).F.E.andB.L.acknowledgefundingbytheAustrian ScienceFundationFWF(projectI5825).b. CONFLICT OF INTEREST STATEMENT Marc Cadotte is the Senior Editor of Ecological Solutions and Evidence,buttooknopartinthepeerreviewanddecision-making processes for this paper. The authors declare no further conflicts of interest. PEER REVIEW The peer review history for this article is available at https:// www. webof scien ce. com/ api/ g atew ay/ wos/ peerreview/ 10. 1002/ 26888319. 70020 . 26888319, 2025, 1, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1002/2688-8319.70020 by Readcube (Labtiva Inc.), Wiley Online Library on [02/05/2025]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License