Non-native tree (NNT) species have been transported worldwide to create or enhance services that are fundamental for human well-being, such as timber provision, erosion control or ornamental value; yet NNTs can also produce undesired effects, such as fire proneness or pollen allergenicity. Despite the variety of effects that NNTs have on multiple ecosystem services, a global quantitative assessment of their costs and benefits is still lacking. Such information is critical for decision-making, management and sustainable exploitation of NNTs. We present here a global assessment of NNT effects on the three main categories of ecosystem services, including regulating (RES), provisioning (PES) and cultural services (CES), and on an ecosystem disservice (EDS), i.e. pollen allergenicity. By searching the scientific literature, country forestry reports, and social media, we compiled a global data set of 1683 case studies from over 125 NNT species, covering 44 countries, all continents but Antarctica, and seven biomes. Using different meta-analysis techniques, we found that, while NNTs increase most RES (e.g. climate regulation, soil erosion control, fertility and formation), they decrease PES (e.g. NNTs contribute less than native trees to global timber provision). Also, they have different effects on CES (e.g. increase aesthetic values but decrease scientific interest), and no effect on the EDS considered. NNT effects on each ecosystem (dis)service showed a strong context dependency, varying across NNT types, biomes and socio-economic conditions. For instance, some RES are increased more by NNTs able to fix atmospheric nitrogen, and when the ecosystem is located in low-latitude biomes; some CES are increased more by NNTs in less-wealthy countries or in countries with higher gross domestic products. The effects of NNTs on several ecosystem (dis)services exhibited some synergies (e.g. among soil fertility, soil formation and climate regulation or between aesthetic values and pollen allergenicity), but also trade-offs (e.g. between fire regulation and soil erosion control). Our analyses provide a quantitative understanding of the complex synergies, trade-offs and context dependencies involved for the effects of NNTs that is essential for attaining a sustained provision of ecosystem services.

Centre for Invasion Biology Department of Botany and Zoology Stellenbosch University Matieland 7602 South Africa

College of Agriculture Polytechnic Institute of Coimbra 3045 601 Coimbra Portugal Centre for Applied Ecology Prof Baeta Neves School of Agriculture University of Lisbon PT1349 017 Lisbon Portugal

College of Agriculture Polytechnic Institute of Coimbra 3045 601 Coimbra Portugal Centre for Functional Ecology Department of Life Sciences University of Coimbra 3000 456 Coimbra Portugal

CREA Research Centre for Foresty and Wood Viale Santa Margherita 80 52100 Arezzo Italy

CSIRO Land and Water Ecosciences Precinct Dutton Park Queensland 4102 Australia

CSIRO Land and Water Wembley Western Australia 6913 Australia School of Biological Sciences University of Western Australia Crawley Western Australia 6009 Australia

Departamento de Biología Facultad de Cc del Mar y Ambientales Instituto Universitario de Investigación Marina Campus de Excelencia Internacional del Mar CEIMAR Universidad de Cádiz E 11510 Puerto Real Spain

Departamento de Ciencias de la Vida Facultad de Ciencias Universidad de Alcalá E 28805 Alcalá de Henares Spain

Department of Biology University of Hawai'i at Hilo Hilo HI 96720 U S A

Department of Botany and Biodiversity Research University of Vienna 1030 Vienna Austria

Department of Ecology and Evolutionary Biology University of California Santa Cruz CA 95060 U S A

Department of Integrative Ecology Estación Biológica de Doñana E 41092 Sevilla Spain

Faculty of Environmental Sciences and Natural Resource Management Norwegian University of Life Sciences Ås Norway

Faculty of Forestry and Wood Technology Mendel University in Brno 613 00 Brno sever Czech Republic Siberian Federal University Krasnoyarsk Krasnoyarsk 660041 Russia

French Associates Institute for Agriculture and Biotechnology of Drylands Blaustein Institutes for Desert Research Ben Gurion University of the Negev Beersheba 84990 Israel

Grupo de Ecología de Invasiones INIBIOMA Universidad Nacional del Comahue CONICET Avenida de los Pioneros 2350 San Carlos de Bariloche Río Negro Argentina

IASMA Research and Innovation Centre Fondazione Edmund Mach 38010 Trento Italy MOUNTFOR Project Centre European Forest Institute 38010 Trento Italy

Landcare Research Lincoln 7640 New Zealand

Natural Resources Institute Finland FI 00791 Helsinki Finland

Research Network in Biodiversity and Evolutionary Biology Research Centre in Biodiversity and Genetic Resources Universidade do Porto PT4485 661 Vairão Portugal Faculdade de Ciências Universidade do Porto PT4169 007 Porto Portugal

Research Network in Biodiversity and Evolutionary Biology Research Centre in Biodiversity and Genetic Resources Universidade do Porto PT4485 661 Vairão Portugal Laboratory of Applied Ecology CITAB Centre for the Research and Technology of Agro Environment and Biological Sciences University of Trás os Montes e Alto Douro Vila Real Portugal

School of Ecosystem and Forest Sciences Faculty of Science The University of Melbourne Richmond Victoria 3121 Australia

Setor de Ecologia Departamento de Biologia Universidade Federal de Lavras Lavras MG 37200 000 Brazil

The University of Queensland School of Biological Sciences Brisbane Queensland 4072 Australia

University of Göttingen 37073 Göttingen Germany

USDA Forest Service Institute of Pacific Islands Forestry Hilo HI U S A

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