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Naturalized Alien Flora of the World: Species Diversity, Taxonomic and Phylogenetic Patterns, Geographic Distribution and Global Hotspots of Plant Invasion

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Naturalized Alien Flora of the World: Species Diversity, Taxonomic and Phylogenetic Patterns, Geographic Distribution and Global Hotspots of Plant Invasion Preslia 89: 203–274, 2017 203 Naturalized alien flora of the world: species diversity, taxonomic and phylogenetic patterns, geographic distribution and global hotspots of plant invasion Naturalizovaná nepůvodní flóra světa: druhová diverzita, taxonomické a fylogenetické složení, geogra- fické zákonitosti a globální ohniska rostlinných invazí Petr P y š e k1,2,3,JanPergl1,Franz Essl4,3,Bernd Lenzner4,Wayne Dawson5, Holger K r e f t6, Patrick W e i g e l t6, Marten Winter7, John K a r t e s z8,Misako Nishino8,LiubovA.Antonova9, Julie F. B a r c e l o n a10,FranciscoJ.Cabezas11, Dairon C á r d e n a s12,JulianaCárdenas-Toro13,14,NicolásCastańo12, Eduardo Chacón4,15, Cyrille Chatelain16, Stefan Dullinger4, Aleksandr L. E b e l17, Estrela Figueiredo18,19,NicolFuentes20, Piero Genovesi21,22, Quentin J. Groom23, Lesley Henderson24, Inderjit25, Andrey K u p r i y a n o v26, Silvana Masciadri27,NoëlieMaurel28,Jan Meerman29,Olga Morozova30, Dietmar Moser4,DanielNickrent31, Pauline M. N o w a k32, Shyama P a g a d33, Annette Patzelt34, Pieter B. P e l s e r10, Hanno Seebens35,Wen-shengShu36, Jacob Thomas37, Mauricio V e l a y o s11, Ewald W e b e r38,JanJ.Wieringa39,40,MaríaP. Baptiste13 &Markvan Kleunen28,41 1Institute of Botany, Department of Invasion Ecology, The Czech Academy of Sciences, CZ-252 43 Průhonice, Czech Republic, e-mail: [email protected], [email protected]; 2Department of Ecology, Faculty of Science, Charles University, CZ-128 44 Viničná 7, Prague 2, Czech Republic; 3Centre for Invasion Biology, Department of Botany & Zool- ogy, Stellenbosch University, Matieland 7602, South Africa; 4Division of Conservation Biology, Vegetation and Landscape Ecology, University of Vienna, 1030 Wien, Austria, e-mail: [email protected], [email protected]; 5DepartmentofBiosci- ences, Durham University, South Road, Durham, DH1 2LF, United Kingdom, e-mail: [email protected]; 6Biodiversity, Macroecology & Biogeography, University of Goettingen, Büsgenweg 1, D-37077 Göttingen, Germany, e-mail: [email protected], [email protected]; 7German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, D-04103 Leipzig, Germany, e-mail: [email protected]; 8Biota of North America Program (BONAP), 9319 Bracken Lane, Chapel Hill, North Carolina, 27516, USA; 9Institute for Aquatic and Ecological Problems, Far East Branch, Russian Academy of Sciences, 680000 Khabarovsk, Russia; 10School of Biological Sciences, University of Canterbury, Private Bag 4800, Christ- church 8140, New Zealand; 11Departamento de Biodiversidad y Conservación, Real Jardín Botánico, CSIC, Plaza de Murillo 2, 28014 Madrid, Spain; 12Instituto Amazónico de Investigaciones Científicas Sinchi, Herbario Amazónico Colombiano, Bogotá, Colombia; 13Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia; 14Arts Faculty, Monash University, Melbourne, Australia; 15Escuela de Biología, Universidad de Costa Rica, 11501 San José, Costa Rica; 16Conservatoire et jardin botaniques de la Ville de Genčve, Genčve, Switzerland; 17Laboratory of Plant doi: 10.23855/preslia.2017.203 204 Preslia 89: 203–274, 2017 Taxonomy and Phylogeny, Tomsk State University, Lenin Prospect 36, 634050 Tomsk, Russia; 18Department of Botany, P.O. Box 77000, Nelson Mandela Metropolitan Univer- sity, Port Elizabeth, 6031 South Africa; 19Centre for Functional Ecology, Departamento de Cięncias da Vida, Universidade de Coimbra, 3001-455 Coimbra, Portugal; 20Departa- mento de Botánica, Facultad de Ciencias Naturales y Oceanográficas, Universidad de Concepción, Casilla 160-C, Concepción, Chile; 21Institute for Environmental Protection and Research (ISPRA), Via Vitaliano Brancati 48, 00144 Rome, Italy; 22Chair IUCN Species Survival Commission’s Invasive Species Specialist Group (ISSG), Rome, Italy; 23Botanic Garden Meise, Domein van Bouchout, B-1860, Meise, Belgium; 24ARC-Plant Protection Research Institute, Pretoria 0001, South Africa; 25Department of Environ- mental Studies, Centre for Environmental Management of Degraded Ecosystems, Uni- versity of Delhi, Delhi 110007, India; 26Institute of Human Ecology, Siberian Branch of Russian Academy of Sciences, Prospect Leningradasky 10, 650065 Kemerovo, Russia; 27Oceanología, Facultad de Ciencias, Universidad de la República, Iguá, 4225, CP 11400, Montevideo, Uruguay; 28Ecology, Department of Biology, University of Konstanz, Universitätsstrasse 10, D-78464 Konstanz, Germany, e-mail: mark.vankleunen@ uni-konstanz.de; 29Belize Tropical Forest Studies, P.O. Box 208, Belmopan, Belize; 30Institute of Geography, Russian Academy of Sciences, Staromonetny 29, 119017 Mos- cow, Russia; 31Department of Plant Biology, Southern Illinois University, Carbondale, Illinois 62901-6509, USA; 32Department of Geography, University Marburg, Deutsch- hausstraße 10, D-35032 Marburg, Germany; 33IUCN Species Survival Commission’s Invasive Species Specialist Group (ISSG), University of Auckland, New Zealand; 34Oman Botanic Garden, Diwan of Royal Court, 122 Muscat, Oman; 35Senckenberg Biodiversity and Climate Research Centre (BiK-F), Senckenberganlage 25, D-60325 Frankfurt am Main, Germany; 36State Key Laboratory of Biocontrol and Guangdong Key Laboratory of Plant Resources, College of Ecology and Evolution, Sun Yat-sen University, Guang- zhou 510275, China; 37Department of Botany & Microbiology, College of Science, King Saud University, P.O. Box 2455, Riyadh 11451, Saudi Arabia; 38Institute of Biochemistry and Biology, University of Potsdam, D-14469 Potsdam, Germany; 39Naturalis Biodiver- sity Center (Botany Section), Darwinweg 2, 2333 CR Leiden, the Netherlands; 40Biosys- tematics Group, Wageningen University, Droevendaalsesteeg 1, 6708 PB Wageningen, the Netherlands; 41Zhejiang Provincial Key Laboratory of Plant Evolutionary Ecology and Conservation, Taizhou University, Taizhou 318000, China Pyšek P., Pergl J., Essl F., Lenzner B., Dawson W., Kreft H., Weigelt P., Winter M., Kartesz J., Nishino M., Antonova L. A., Barcelona J. F., Cabezas F. J., Cárdenas D., Cárdenas-Toro J., Castańo N., Chacón E., Chatelain C., Dullinger S., Ebel A. L., Figueiredo E., Fuentes N., Genovesi P., Groom Q. J., Henderson L., Inderjit, Kupriyanov A., Masciadri S., Maurel N., Meerman J., Morozova O., Moser D., Nickrent D., Nowak P. M., Pagad S., Patzelt A., Pelser P. B., Seebens H., Shu W., Thomas J., Velayos M., Weber E., Wieringa J. J., Baptiste M. P. & van Kleunen M. (2017): Naturalized alien flora of the world: species diversity, taxonomic and phylogenetic patterns, geographic distribution and global hotspots of plant invasion. – Preslia 89: 203–274. Using the recently built Global Naturalized Alien Flora (GloNAF) database, containing data on the distribution of naturalized alien plants in 483 mainland and 361 island regions of the world, we describe patterns in diversity and geographic distribution of naturalized and invasive plant species, taxonomic, phylogenetic and life-history structure of the global naturalized flora as well Pyšek et al.: Naturalized alien flora of the world 205 as levels of naturalization and their determinants. The mainland regions with the highest numbers of naturalized aliens are some Australian states (with New South Wales being the richest on this continent) and several North American regions (of which California with 1753 naturalized plant species represents the world’s richest region in terms of naturalized alien vascular plants). Eng- land, Japan, New Zealand and the Hawaiian archipelago harbour most naturalized plants among islands or island groups. These regions also form the main hotspots of the regional levels of natu- ralization, measured as the percentage of naturalized aliens in the total flora of the region. Such hotspots of relative naturalized species richness appear on both the western and eastern coasts of North America, in north-western Europe, South Africa, south-eastern Australia, New Zealand, and India. High levels of island invasions by naturalized plants are concentrated in the Pacific, but also occur on individual islands across all oceans. The numbers of naturalized species are closely correlated with those of native species, with a stronger correlation and steeper increase for islands than mainland regions, indicating a greater vulnerability of islands to invasion by species that become successfully naturalized. South Africa, India, California, Cuba, Florida, Queensland and Japan have the highest numbers of invasive species. Regions in temperate and tropical zonobiomes harbour in total 9036 and 6774 naturalized species, respectively, followed by 3280 species naturalized in the Mediterranean zonobiome, 3057 in the subtropical zonobiome and 321 in the Arctic. The New World is richer in naturalized alien plants, with 9905 species compared to 7923 recorded in the Old World. While isolation is the key factor driving the level of naturaliza- tion on islands, zonobiomes differing in climatic regimes, and socioeconomy represented by per capita GDP, are central for mainland regions. The 11 most widely distributed species each occur in regions covering about one third of the globe or more in terms of the number of regions where they are naturalized and at least 35% of the Earth’s land surface in terms of those regions’ areas, with the most widely distributed species Sonchus oleraceus occuring in 48% of the regions that cover 42% of the world area. Other widely distributed species are Ricinus communis, Oxalis corniculata, Portulaca oleracea, Eleusine indica, Chenopodium album,
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