Naturalized alien flora of the world: species diversity, taxonomic and phylogenetic patterns, geographic distribution and global hotspots of plant invasion

Petr Pyšek 1 2 3 , Jan Pergl 1 , Franz Essl 3 4 , Bernd Lenzner 4 , Wayne Dawson 5 , Holger Kreft 6 , Patrick Weigelt 6 , Marten Winter 7 , John Kartesz 8 , Misako Nishino 8 , Liubov A. Antonova 9 , Julie F. Barcelona 10 , Francisco J. Cabezas 11 , Dairon Cárdenas 12 , Juliana Cárdenas-Toro 13 14 , Nicolás Castańo 12 , Eduardo Chacón 4 15 , Cyrille Chatelain 16 , Stefan Dullinger 4 , Aleksandr L. Ebel 17 , Estrela Figueiredo 18 19 , Nicol Fuentes 20 , Piero Genovesi 21 22 , Quentin J. Groom 23 , Lesley Henderson 24 , Inderjit 25 , Andrey Kupriyanov 26 , Silvana Masciadri 27 , Noëlie Maurel 28 , Jan Meerman 29 , Olga Morozova 30 , Dietmar Moser 4 , Daniel Nickrent 31 , Pauline M. Nowak 32 , Shyama Pagad 33 , Annette Patzelt 34 , Pieter B. Pelser 10 , Hanno Seebens 35 , Wen-sheng Shu 36 , Jacob Thomas 37 , Mauricio Velayos 11 , Ewald Weber 38 , Jan J. Wieringa 39 40 , María P. Baptiste 13 & Mark van Kleunen 28 41


  1. Institute of Botany, Department of Invasion Ecology, The Czech Academy of Sciences, CZ-252 43 Průhonice, Czech Republic
  2. Department of Ecology, Faculty of Science, Charles University, CZ-128 44 Viničná 7, Prague 2, Czech Republic
  3. Centre for Invasion Biology, Department of Botany & Zoology, Stellenbosch University, Matieland 7602, South Africa
  4. Division of Conservation Biology, Vegetation and Landscape Ecology, University of Vienna, 1030 Wien, Austria
  5. Department of Biosciences, Durham University, South Road, Durham, DH1 2LF, United Kingdom
  6. Biodiversity, Macroecology & Biogeography, University of Goettingen, Büsgenweg 1, D-37077 Göttingen, Germany
  7. German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, D-04103 Leipzig, Germany
  8. Biota of North America Program (BONAP), 9319 Bracken Lane, Chapel Hill, North Carolina, 27516, USA
  9. Institute for Aquatic and Ecological Problems, Far East Branch, Russian Academy of Sciences, 680000 Khabarovsk, Russia
  10. School of Biological Sciences, University of Canterbury, Private Bag 4800, Christchurch 8140, New Zealand
  11. Departamento de Biodiversidad y Conservación, Real Jardín Botánico, CSIC, Plaza de Murillo 2, 28014 Madrid, Spain
  12. Instituto Amazónico de Investigaciones Científicas Sinchi, Herbario Amazónico Colombiano, Bogotá, Colombia
  13. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia
  14. Arts Faculty, Monash University, Melbourne, Australia
  15. Escuela de Biología, Universidad de Costa Rica, 11501 San José, Costa Rica
  16. Conservatoire et jardin botaniques de la Ville de Genčve, Genčve, Switzerland
  17. Laboratory of Plant Taxonomy and Phylogeny, Tomsk State University, Lenin Prospect 36, 634050 Tomsk, Russia
  18. Department of Botany, P.O. Box 77000, Nelson Mandela Metropolitan University, Port Elizabeth, 6031 South Africa
  19. Centre for Functional Ecology, Departamento de Cięncias da Vida, Universidade de Coimbra, 3001-455 Coimbra, Portugal
  20. Departamento de Botánica, Facultad de Ciencias Naturales y Oceanográficas, Universidad de Concepción, Casilla 160-C, Concepción, Chile
  21. Institute for Environmental Protection and Research (ISPRA), Via Vitaliano Brancati 48, 00144 Rome, Italy
  22. Chair IUCN Species Survival Commission’s Invasive Species Specialist Group (ISSG), Rome, Italy
  23. Botanic Garden Meise, Domein van Bouchout, B-1860, Meise, Belgium
  24. ARC-Plant Protection Research Institute, Pretoria 0001, South Africa
  25. Department of Environmental Studies, Centre for Environmental Management of Degraded Ecosystems, University of Delhi, Delhi 110007, India
  26. Institute of Human Ecology, Siberian Branch of Russian Academy of Sciences, Prospect Leningradasky 10, 650065 Kemerovo, Russia
  27. Oceanología, Facultad de Ciencias, Universidad de la República, Iguá, 4225, CP 11400, Montevideo, Uruguay
  28. Ecology, Department of Biology, University of Konstanz, Universitätsstrasse 10, D-78464 Konstanz, Germany
  29. Belize Tropical Forest Studies, P.O. Box 208, Belmopan, Belize
  30. Institute of Geography, Russian Academy of Sciences, Staromonetny 29, 119017 Moscow, Russia
  31. Department of Plant Biology, Southern Illinois University, Carbondale, Illinois 62901-6509, USA
  32. Department of Geography, University Marburg, Deutschhausstraße 10, D-35032 Marburg, Germany
  33. IUCN Species Survival Commission’s Invasive Species Specialist Group (ISSG), University of Auckland, New Zealand
  34. Oman Botanic Garden, Diwan of Royal Court, 122 Muscat, Oman
  35. Senckenberg Biodiversity and Climate Research Centre (BiK-F), Senckenberganlage 25, D-60325 Frankfurt am Main, Germany
  36. State Key Laboratory of Biocontrol and Guangdong Key Laboratory of Plant Resources, College of Ecology and Evolution, Sun Yat-sen University, Guangzhou 510275, China
  37. Department of Botany & Microbiology, College of Science, King Saud University, P.O. Box 2455, Riyadh 11451, Saudi Arabia
  38. Institute of Biochemistry and Biology, University of Potsdam, D-14469 Potsdam, Germany
  39. Naturalis Biodiversity Center (Botany Section), Darwinweg 2, 2333 CR Leiden, the Netherlands
  40. Biosystematics Group, Wageningen University, Droevendaalsesteeg 1, 6708 PB Wageningen, the Netherlands
  41. Zhejiang Provincial Key Laboratory of Plant Evolutionary Ecology and Conservation, Taizhou University, Taizhou 318000, China

Published: 12 July 2017 ,

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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 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). England, 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 naturalization, 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 naturalization 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, Capsella bursa-pastoris, Stellaria media, Bidens pilosa, Datura stramonium and Echinochloa crus-galli. Using the occurrence as invasive rather than only naturalized yields a different ranking, with Lantana camara (120 regions out of 349 for which data on invasive status are known), Calotropis procera (118), Eichhornia crassipes (113), Sonchus oleraceus (108) and Leucaena leucocephala (103) on top. As to the life-history spectra, islands harbour more naturalized woody species (34.4%) thanmainland regions (29.5%), and fewer annual herbs (18.7% compared to 22.3%). Ranking families by their absolute numbers of naturalized species reveals that Compositae (1343 species), Poaceae (1267) and Leguminosae (1189) contribute most to the global naturalized alien flora. Some families are disproportionally represented by naturalized aliens on islands (Arecaceae, Araceae, Acanthaceae, Amaryllidaceae, Asparagaceae, Convolvulaceae, Rubiaceae, Malvaceae), and much fewer so on mainland (e.g. Brassicaceae, Caryophyllaceae, Boraginaceae). Relating the numbers of naturalized species in a family to its total global richness shows that some of the large species-rich families are over-represented among naturalized aliens (e.g. Poaceae, Leguminosae, Rosaceae, Amaranthaceae, Pinaceae), some under-represented (e.g. Euphorbiaceae, Rubiaceae), whereas the one richest in naturalized species, Compositae, reaches a value expected from its global species richness. Significant phylogenetic signal indicates that families with an increased potential of their species to naturalize are not distributed randomly on the evolutionary tree. Solanum (112 species), Euphorbia (108) and Carex (106) are the genera richest in terms of naturalized species; over-represented on islands are Cotoneaster, Juncus, Eucalyptus, Salix, Hypericum, Geranium and Persicaria, while those relatively richer in naturalized species on the mainland are Atriplex, Opuntia, Oenothera, Artemisia, Vicia, Galium and Rosa. The data presented in this paper also point to where information is lacking and set priorities for future data collection. The GloNAF database has potential for designing concerted action to fill such data gaps, and provide a basis for allocating resources most efficiently towards better understanding and management of plant invasions worldwide.


alien species, distribution, Global Naturalized Alien Flora (GloNAF) database, invasive species, islands, life history, mainland, naturalized species, phylogeny, plant invasion, regional floras, species richness, taxonomy, zonobiome

How to cite

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: 203274,