Ecol Solut Evid. 2025;6:e70020. | 1 of 11 https://doi.org/10.1002/2688-8319.70020 wileyonlinelibrary.com/journal/eso3 Received:12December2024 | Accepted:25January2025 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 CreativeCommonsAttribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. ©2025TheAuthor(s).Ecological Solutions and EvidencepublishedbyJohnWiley&SonsLtdonbehalfofBritishEcologicalSociety. DaijiangLiandLukeJ.Potgietercontributedequallytothiswork. For affiliations refer to page 9. Correspondence Daijiang Li Email:[email protected] LukeJ.Potgieter Email:
[email protected] MarcW.Cadotte Email:
[email protected] Funding information NaturalSciencesandEngineering Research Council of Canada, Grant/AwardNumber:#RGPIN- 202203579 and 386151; Deutsche Forschungsgemeinschaft, Grant/ AwardNumber:264740629,FZT118 and202548816;NationalScience Abstract 1. Urban areas are foci for the introduction of nonnative plant species, and they oftenactaslaunchingsitesforinvasionsintothewiderenvironment.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. WeassembleddatafrommultiplesourcestocreatetheGlobalUrbanBiological InvasionsCompendium(GUBIC)forvascularplantsrepresenting553urbancentresfrom61countriesacrosseverycontinentexceptAntarctica. 3. TheGUBICrepositoryincludes8140non-nativeplantspeciesfrom253families. 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 densityandextensivelandscapemodification,presentuniqueopportunities 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(Gallardoetal.,2016;Potgieteretal.,2024).Urbanplant 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;Potgieteretal.,2017).However,thereisalackof 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 mightprovide(Milanovićetal.,2020),withcurrentknowledgeremaining geographically heterogeneous and focused on only a few well-studiedtaxa(Vazetal.,2018). Frameworks for understanding and managing urban plant invasionsarelessfrequentlystudiedthaninotherhabitats(but see Gaertner et al., 2016;Potgieter&Cadotte,2020).Whileexisting 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.Managersinurbanareasfaceuniquechallengesduetothe interplay between the built environment and complex socioeconomic factors, which can significantly alter ecosystem conditions. However,thesechallengeshaveonlyrecentlybeenincorporated into models to predict urban invasion dynamics and impacts and identifyappropriatemanagementstrategies(Gaertneretal.,2016; Potgieteretal.,2022). Despite these advances, empirical studies on urban biological invasions remain limited, particularly in terms of taxonomic coverageandspatialscale(Cadotteetal.,2017).Mostempirical 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. Althoughnumerousregionalandcity-specificinventoriesofnon- native species exist, these are often from uncoordinated efforts carried out independently by research groups focusing on particularresearchquestions.Therefore,thesediverseresourceslack 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 greyliteratureandonlyavailableinnon-Englishlanguages,limiting theiraccessibility.Whilethesebiologicalinventoriesarecrucial 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 isrequiredtounderstandtheroleofurbanareasinshapingthe patternsofplant invasionsandtheunderlying 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 Tocompilealistofnon-nativeplantspeciesinurbanareas(see Section2.1.2 for the methods used to delineate urban boundaries)globally,wecombinedmultipledatasources.Thisapproach 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, invasionbiology,macroecology,conservation,urbanplanningandsustainability.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 Alienspecies,biodiversitychange,biologicalinvasions,cities,naturalizedspecies,non-native plants, urbanization Foundation,Grant/AwardNumber: DEB-2213567;TheItalianMinistryof UniversityandResearch,Grant/Award Number:CN00000033;CNPq-Brazil, Grant/AwardNumber:302643/2022-2; CzechScienceFoundation,Grant/Award Number:19-28491Xand25-15190S; CzechAcademyofSciences,Grant/Award Number:RVO67985939 Handling Editor:HollyJones 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. Weincludedonlyestablishednon-nativeplantspecies,whichare those with selfsustaining populations, also commonly referred to asnaturalized(Blackburnetal.,2011; Richardson et al., 2000; see Section2.1.5). 2.1 | Data acquisition and compilation 2.1.1 | Datasource1:GlobalUrbanBiological Invasions Consortium An international workshop to address biological invasions in urbanecosystemswashostedbytheCentreforInvasionBiology in Stellenbosch, South Africa, in November 2016 (Gaertner et al., 2017).ThisworkshopledtothecreationoftheGlobal Urban BiologicalInvasionsConsortium, which hosted a coordinating meetinginJune2019thatbroughttogethermorethan70researchers from14countriesfromallcontinentsexceptAntarctica.Oneofthe prioritized activities was to compile lists of nonnative plant speciesforurbanareas.Aworkinggroup“SynthesizingGlobalUrban BiologicalInvasionKnowledge”(sGUBIK,fundedbysDiv,thesynthesiscentreofiDiv,theGermanCentreforIntegrativeBiodiversity 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. Wecompileddatausingthefollowing approaches.First,we sentarequesttoover150membersoftheGlobalUrbanBiological Invasions Consortium in 2019 to upload datasets for any urban taxatoaSharePointrepositoryattheUniversityofToronto.The cut-offforthedatarequestwasDecember2021.Second,during AugusttoNovember2019,wesearchedthepublishedliteraturein English,PortugueseandSpanishaswellastheDryaddatarepository(www. datad ryad. org)forstudiesanddatasetscontainingspecies 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 includedtheUrbanBiodiversityResearchCoordinationNetwork (UrBioNet) dataset, a large multi-city compilation (Aronson et al., 2014),featuring14,240spontaneousplantspecies(i.e.not cultivatedorplanted),ofwhich4241areidentifiedasnon-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 urbancentreswithinthesamecountrysharedthesamename(e.g. Madison,Wisconsinvs.Madison,IndianaintheUnitedStates),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. Asaresult,datasetscollectedfromdatacontributors,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 | Datasource2:GlobalBiodiversity Information Facility BeforeextractingoccurrencedataforeachurbanareafromtheGlobal Biodiversity Information Facility (GBIF), we delineated the boundariesofurbanareas.Weusedtheglobalurbancentresdataprovided bytheGlobalHumanSettlementLayer(GHSL,Pesaresietal.,2019, https:// ghsl. jrc. ec. europa. eu/ ucdb2 018Ov erview. php), which definesurbancentresascontiguous1 km2 grid cells with a population density of at least 1500 inhabitants per km2ofpermanentland(areas that are consistently above water and exclude bodies of water, such asoceans,seas,largeriversandlakes)orwithmorethan50%built-up surface shared on permanent land and with at least 50,000 inhabitantsintheclusterwithsmoothedboundariesandsmallgaps(<15 km2) filled. Overall, there are 13,189 unique urban centres worldwide. Subsequently,smaller,nearbyurbancentreslocatedwithina5 kmradius 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.Werefrainedfromfurthermergingsmallercentresthat,although withina5 kmradiusofthepreviouslymergedsmallercentres,were situatedbeyondthe5 kmboundaryfromthelargerurbancentre.This processresultedin11,621uniqueurbancentresglobally. InAugust2023,wequeriedGBIFanddownloadedplantoccurrence records from each urban centre to compile the flora of these urbanareas(seeTable S1fortheDOIsofdownloadeddatasets).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’(Zizkaetal.,2019).Wealsoremovedallrecords with identification above species level, fossil specimens, preserved specimens, living specimens and those with locality uncertainty greaterthan30 kmorwithina500 mvicinityofbiodiversityinstitutions,botanicgardens,zoos,museums,GBIFheadquarters,etc. 2.1.3 | Qualitycontrolandmergingofdata Beforemergingdatafromsources1and2,weconductedpreliminary filteringof thesedatasets.ForeachurbancentrewithGBIF data, we used the number of observations of each species as a proxy fortheabundanceofthatspecies.Wecalculatedobservedspecies richnessandestimatedspeciesrichnessusingtheChao1equation, whichincorporatessingletonsanddoubletons(i.e.speciesobserved onlyonceortwice): (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 centreusingtheRpackage‘iNEXT’forrarefaction(Chaoetal.,2014; Hsiehetal.,2024). WeconsideredanurbancentretohaverobustGBIFdataif:(1)it hadover1000observedplantspecies;(2)thecommunitysamplecoverage was >90%;and(3)theobservedspeciesrichnesswasgreater than75%oftheestimatedspeciesrichness.Weusedthesecriteria tobalancethenumberofretainedurbancentresanddataquality.For data source 1, if an urban centre had more than 300 plant species, we retaineditandfurtherintegrateditwithdatasource2(GBIFdata)of thaturbancentreregardlessoftheGBIFdataquality.Ifanurbancentre had fewer than 300 species from data source 1 and did not have adequateGBIFdatacoverage,weremovedthaturbancentrefromour database. If an urban centre had fewer than 300 species from data source 1but hadadequateGBIF data coverage (i.e. met the above threecriteria),weretainedbothdatasourcesforthaturbancentre.We removedthoseurbancentreswithonlyGBIFdatathatdidnotmeet thethreecriteriaabove(seeFigure 1foraschematicworkflow).Like thecriteriaweusedfortheGBIFdata,weselected300specieshere tobalancethenumberofurbancentresandtheirdataqualityafter carefully explored our datasets. The final database included 553 urban centres(Figure 2).Foreachoftheseurbancentres,wederivedalistof established nonnative plant species using the merged data sources. 2.1.4 | Standardizespeciesnames We standardized species and family names against the World Checklist of Vascular Plants (WCVP, Govaerts, 2024) for the mergeddatabaseusingtheRpackagerWCVP(version1.0.3,Brown et al., 2023).WeselectedWCVPasitrepresentsoneofthemostcomprehensive and up-to-date taxonomic resources available (Grenié et al., 2022).WCVPalsoservesasthetaxonomicbackboneforthe mostrecentversionoftheGlobalNaturalizedAlienFlora(GloNAF), whichwasupdatedfollowingvanKleunenetal.(2019).GloNAFwas used to determine whether a species is nonnative in a particular regionwhereanurbancentrewaslocated(seeSection2.1.5below). Notethatspecieswith“unplacednames”(n = 65acrossallspecies) orhasnotmatchfromWCVPwereexcludedfromthefinaldataset (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.Wealsomergedsubspeciesorvarietiestothemainspecies and only kept binomial species names in the final database. 2.1.5 | Cross-validationtodeterminethestatusof species Todistinguishbetweenestablished(naturalized)andnativeorcasual species(i.e.thosethatmightflourishandevenreproduceoccasionally in an area but which do not form selfreplacing populations; FIGURE 1 SchematicfigureshowingtheworkflowofthecompilationoftheGlobalUrbanBiologicalInvasionsCompendiumdatabase. UrBioNet:TheUrbanBiodiversityResearchCoordinationNetwork.GHSL,TheGlobalHumanSettlementLayer;GBIF,TheGlobal BiodiversityInformationFacility;WCVP,TheWorldChecklistofVascularPlants;GloNAF,GlobalNaturalizedAlienFlora. 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 Geographicdistributionofurbancentresacrosstheworld(panela;n = 553)andEurope(panelb)andthenumberof establishednon-nativeplantspeciestheycontain(colouredpoints). # of non−native species (log10) 1.52.0 2.53.0 (a) (b) FIGURE 3 Thedistributionoffamilysizesforthe253establishednon-nativeplantfamiliesinthedataset.Themainplotcontainsthetop 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)locatedinaspecificurbancentre,weused theGloNAFdatabaseasitprovidesthemostupdatedinformation of naturalized plant species across the world. For each urban centre, weusedthedelineatedboundariesprovidedbytheGHSL.Foreach species listed within an urban centre, we classified the species as nonnative to that urban centre if its polygon intersected with the species'naturalizedorinvasiverange.Wealsocross-referencedall species with local checklists of nonnative plant species validated by experts(Kalusováetal.,2024).Therefore,forthoseurbancentres (mostlyinEurope),thelistsofnaturalizedspecieswereslightlydifferentfromthosebasedonGloNAFalone. 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 EuropeanandNorthAmericanurbancentres,whichtogetheraccount 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 253families(Figure 3).Mostfamiliescontainfewspecies,with73 familieseachcontaining20ormorenon-nativespecies(Figure 3). Asteraceae,Poaceae,FabaceaeandRosaceaecontainaboutone- thirdofallspecies(n = 2641;Table 1).Themostwidespreadnon- 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.Ararefactionof speciesoccurrencesacrossurbancentres(Figure 4)showsthatwe are approaching an asymptote with our sample of 553 urban centres.However,thesamplingcurvealsosuggeststhatmoreurban floral sampling is needed, especially from regions with sparse data (e.g.SouthAsia,northernSouthAmerica). TABLE 1 Themostwidespread(top30)establishednon-nativeplantspeciesinurbancentres(n = 553)acrosstheworld.Notethatthislist wasderivedfromdifferentsamplingeffortsandhasabiasinfavourofnon-nativespeciesinEuropeanandNorthAmericanurbancentres. 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 decisionsandrecognizethatlimitationsareinevitable.Someissues to be cognizant of for analysis and interpretation include: 1. Our definition of urbanized areas delineated contiguous areas. Becauseofthisdefinition,someurbanizedareasspanmultiple 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 Thetop20urbancentres(left)andthetop20countries(right)withthegreatestnumberofnon-nativeplantspecies.Notethat theselistsareskewedtowardsEuropeanandNorthAmericanurbancentres(seeFigure 2).Thenumberspresentedforsomecountries (e.g.France)alsoincludednon-nativeplantspeciesfromtheiroverseasurbancentres. Urban centre Number of established nonnative species Country Number of established nonnative species New York 1663 UnitedStatesofAmerica 4409 LosAngeles 1534 Australia 2596 Sydney 1486 France 2187 Philadelphia 1455 NewZealand 1561 Melbourne 1450 Canada 1476 WashingtonD.C. 1414 Russia 1251 Auckland 1310 Japan 1154 Boston 1300 Mexico 1142 SanJose(USA) 1231 Germany 1123 Tijuana 1066 UnitedKingdom 986 St.Louis 1058 Switzerland 966 Tokyo 1038 SouthAfrica 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 Rarefactioncurveofthe 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,notonlyincludesneighbouringcitiesinMichigan, like Dearborn, but also the Canadian city of Windsor. 2. Whilemostrecordedspeciesinourdatasetcanbeconfirmedas 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 naturallyoccurringrecords.Asaresult,thedatashouldbeinterpreted cautiously, particularly when comparing nonnative species richness at broader spatial scales, such as across countries, ratherthanatthecitylevel. Speciesin our dataset withwidespread occurrences across multiple urban centres are likely to be established, whereas species recorded in only one urban centre mightrequirefurtherscrutiny.Werecommendthatusersconsider including these singleton records in sensitivity analyses to assess the robustness of their results. Therefore, the numbers of non-nativespeciesreportedhere(e.g.Table 2)areinsomecases higher than those reported for individual countries in recent studies(Kalusováetal.,2024;Pyšeketal.,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 duringdirectsampling.Becauseourgoalistocompileanon-native flora of urban centres, these limitations do not significantly affect our dataset. 4. ThedataextractedfromGBIFincludegeographicallybiasedand 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),whicharelikelyunderestimates.Analyses of richness and diversity should include rarefaction or some otherwayofaccountingforunequalsamplingasthenumberof nonnative species increased with the number of observations (Figure 5b).Notably,manyurbancentresfromtheGlobalSouth (e.g.India;Figure 2)wereabsentfromourdatabaseduetothe paucity of available data. 5 | CONCLUSION Thedatabasepresentedhererepresentsauniqueandvaluableresource for addressing a wide range of basic and applied ecological questions,particularlythoserelatedtobiologicalinvasions.Thisresource can support hypothesis testing at the macroand global scale (e.g.bioticresistanceorinvasiondebt).Itcanalsobeusedtomodel 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 wasfurtherdevelopedbythesGUBIKworkinggroup(DaijiangLi,Luke J. Potgieter, Myla F. J. Aronson, Irena Axmanová, Benjamin Baiser, MartaCarboni,LauraCelesti-Grapow,SonjaKnapp,IngolfKühn,Ana Carolina Lacerda de Matos, Zdeňka Lososová, Flavia A. Montaño- Centellas,PetrPyšek,DavidM.Richardson,LaurenB.Trotta,RafaelD. ZenniandMarcW.Cadotte).LukeJ.Potgieterledthedatacollection FIGURE 5 (a)Distributionofthenumberofnon-nativeplantspeciesinurbancentres(n = 553)across61countriesand(b)therelationship 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.ThesGUBIKteamsupporteddataintegrationandstandardization.LukeJ.PotgieterandDaijiangLidraftedthemanuscript,withall coauthors contributing to its editing and revision. AFFILIATIONS 1DepartmentofEcologyandEvolutionaryBiology,UniversityofArizona, Tucson,Arizona,USA;2DepartmentofBiologicalSciences,Universityof Toronto-Scarborough,Toronto,Ontario,Canada;3Centre for Invasion Biology,DepartmentofBotany&Zoology,StellenboschUniversity, Stellenbosch,SouthAfrica;4DepartmentofEcology,Evolutionand NaturalResources,Rutgers,TheStateUniversityofNewJersey,New Brunswick,NewJersey,USA;5DepartmentofBotanyandZoology,Faculty ofScience,MasarykUniversity,Brno,CzechRepublic;6Department of WildlifeEcologyandConservation,UniversityofFlorida,Gainesville, Florida,USA;7DepartmentofSciences,RomaTreUniversity,Rome,Italy; 8DepartmentofEnvironmentalBiology,SapienzaUniversityofRome, Rome, Italy; 9DepartmentofCommunityEcology,HelmholtzCentrefor EnvironmentalResearch-UFZ,Halle(Saale),Germany;10German Centre forIntegrativeBiodiversityResearch(iDiv),Halle-Jena-Leipzig,Leipzig, Germany; 11GeobotanyandBotanicalGarden,MartinLutherUniversity Halle-Wittenberg,Halle(Saale),Germany;12DepartamentodeEcologiae Conservação, Instituto de Ciências Naturais, Federal University of Lavras, Lavras,MinasGerais,Brazil;13DepartmentofBiologicalSciences,Louisiana StateUniversity,BatonRouge,Louisiana,USA;14Department of Invasion Ecology,InstituteofBotany,CzechAcademyofSciences,Průhonice, Czech Republic; 15DepartmentofEcology,FacultyofScience,Charles University,Prague,CzechRepublic;16UnitforEnvironmentalSciences andManagement,North-WestUniversity,Potchefstroom,SouthAfrica; 17EnvironmentalResearchInstitute,UniversityofWaikato,Hamilton,New Zealand;18EcologyGroup,DepartmentofBiology,UniversityofKonstanz, Konstanz,Germany;19FriesnerHerbarium,ButlerUniversity,Indianapolis, Indiana,USA;20InstituteofDendrology,PolishAcademyofSciences, Kórnik,Poland;21DivisionofBioInvasions,GlobalChange&Macroecology, DepartmentofBotanyandBiodiversityResearch,UniversityofVienna, Vienna,Austria;22DepartmentofEcologyandEvolutionaryBiology, UniversityofTennessee,Knoxville,Tennessee,USA;23Biodiversityand Landscape,TERRAResearchCentre,GemblouxAgro-BioTech,Universityof Liège,Gembloux,Belgium;24PlantsandEcosystems,UniversityofAntwerp, Wilrijk,Belgium;25Nürtingen-GeislingenUniversity(HfWU),Nürtingen, Germany; 26ThünenInstitute,InstituteofBiodiversity,Braunschweig, Germany; 27CentreforInvasionBiology,DepartmentofMathematical Sciences,StellenboschUniversity,Stellenbosch,SouthAfrica;28African InstituteforMathematicalSciences,NationalInstituteforNational InstituteforTheoreticalandComputationalSciences,CapeTown,South Africa;29CentreforBiodiversity&Taxonomy,DepartmentofBotany, UniversityofKashmir,Srinagar,JammuandKashmir,India;30Department ofEcologyandEvolutionaryBiology,UniversityofTorontoMississauga, Mississauga, Ontario, Canada; 31Biodiversity,Macroecology&Biogeography, University of Göttingen, Göttingen, Germany; 32CentreofBiodiversityand SustainableLandUse(CBL),UniversityofGöttingen,Göttingen,Germany; 33DepartmentofEcologyandEvolutionaryBiology,YaleUniversity,New Haven,Connecticut,USA;34CenterforBiodiversityandGlobalChange, YaleUniversity,NewHaven,Connecticut,USA;35Ecology&Biodiversity, DepartmentofBiology,UtrechtUniversity,Utrecht,TheNetherlands; 36CollegeofForestry,WildlifeandEnvironment,AuburnUniversity,Auburn, Alabama,USA;37CentrodeInvestigaciónenBiodiversidadyConservación, UniversidadAutónomadelEstadodeMorelos,Cuernavaca,Mexico; 38ResearchCenterforAdvancedScienceandTechnology,TheUniversityof Tokyo,Tokyo,Japan;39SchoolofNaturalResources,UniversityofMissouri, Columbia,Missouri,USA;40Ekaterinburg,Russia;41FacultyofEnvironmental Studies,TokyoCityUniversity,Tokyo,Japan;42ZhejiangProvincialKey LaboratoryofPlantEvolutionaryEcologyandConservation,Taizhou University, Taizhou, China; 43DepartamentodeBiologíaVegetalyEcología, UniversidaddeSevilla,Sevilla,Spain;44EstaciónBiológicadeDoñana (EBD-CSIC),Sevilla,Spain;45KeyLaboratoryofTropicalBiologicalResources ofMinistryofEducation,SchoolofLifeandPharmaceuticalSciences,Hainan University,Haikou,China;46DepartmentofEnvironmentalScience,Radboud InstituteforBiologicalandEnvironmentalSciences,RadboudUniversity, Nijmegen, The Netherlands; 47InstituteforHousingandEnvironment, Darmstadt, Germany and 48SchoolofAgricultureFoodandEcosystem Science,UniversityofMelbourne,Parkville,Victoria,Australia FUNDING INFORMATION This paper is a joint effort of the working group sGUBIK kindly supportedbysDiv,theSynthesisCentreoftheGermanCentrefor IntegrativeBiodiversityResearch(iDiv)Halle-Jena-Leipzig,funded by the German Research Foundation (FZT 118, 202548816). The Global Urban Biological Invasion Consortium was initially funded bytheConnaughtGlobalChallengesAward,theOfficeoftheVice- PresidentInternational,theSchoolofGraduateStudies,University of Toronto, and the Office of the Vice-Principal Research at the UniversityofTorontoScarborough.L.J.P.andD.M.R.acknowledge support from the Centre for Invasion Biology and Stellenbosch University.D.L.wassupportedby USNSFDEB-2213567.M.W.C. wassupportedbytheNaturalSciencesandEngineeringResearch CouncilofCanada(#386151).Z.L.andP.P.weresupportedbygrant no.25-15190S(CzechScienceFoundation)andlong-termresearch developmentprojectRVO67985939(CzechAcademyofSciences). Z.L.andI.A.weresupportedbyEXPROgrantno.19-28491X(Czech Science Foundation). M.C. acknowledges the support of NBFC, fundedbytheItalianMinistryofUniversityandResearch,PNRR, Missione4Componente2,“Dallaricercaall'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 supportedby the Natural Sciences andEngineering ResearchCouncil ofCanada(Discovery Grant #RGPIN-2022-03579). M.V. by (PID2021-122690OB-I00) funded by MCIN/AEI/10.13039/501100011033/FEDER, UE. G.H. acknowledgesthesupportofAlexandervonHumboldtFoundation andCoordenaçãodeAperfeiçoamentodePessoaldeNívelSuperior– Brasil(Capes)–Financecode001.M.K.D.acknowledgesthesupport oftheInstituteofDendrology,PolishAcademyofSciences.A.S.M wassupportedbytheMitsui&Co.,Ltd.EnvironmentFund(funding no.R17-0062).F.E.andB.L.acknowledgefundingbytheAustrian ScienceFundationFWF(projectI5825).b. CONFLICT OF INTEREST STATEMENT Marc Cadotte is the Senior Editor of Ecological Solutions and Evidence,buttooknopartinthepeerreviewanddecision-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