RT Journal Article T1 Global impoverishment of natural vegetation revealed by dark diversity A1 Pärtel, Meelis A1 Tamme, Riin A1 Carmona, Carlos P. A1 Riibak, Kersti A1 Moora, Mari A1 Bennett, Jonathan A. A1 Chiarucci, Alessandro A1 Chytrý, Milan A1 de Bello, Francesco A1 Eriksson, Ove A1 Harrison, Susan A1 Lewis, Robert John A1 Moles, Angela T. A1 Öpik, Maarja A1 Price, Jodi N. A1 Amputu, Vistorina A1 Askarizadeh, Diana A1 Atashgahi, Zohreh A1 Aubin, Isabelle A1 Azcárate, Francisco M. A1 Barrett, Matthew D. A1 Bashirzadeh, Maral A1 Bátori, Zoltán A1 Beenaerts, Natalie A1 Bergholz, Kolja A1 Birkeli, Kristine A1 Biurrun, Idoia A1 Blanco-Moreno, José M. A1 Bloodworth, Kathryn J. A1 Boisvert-Marsh, Laura A1 Boldgiv, Bazartseren A1 Brancalion, Pedro H. S. A1 Brearley, Francis Q. A1 Brown, Charlotte A1 Bueno, C. Guillermo A1 Buffa, Gabriella A1 Cahill, James F. A1 Campos, Juan A. A1 Cangelmi, Giacomo A1 Carbognani, Michele A1 Carcaillet, Christopher A1 Cerabolini, Bruno E. L. A1 Chevalier, Richard A1 Clavel, Jan S. A1 Costa, José M. A1 Cousins, Sara A. O. A1 Čuda, Jan A1 Dairel, Mariana A1 Dalle Fratte, Michele A1 Danilova, Alena A1 Davison, John A1 Deák, Balázs A1 Del Vecchio, Silvia A1 Dembicz, Iwona A1 Dengler, Jürgen A1 Dolezal, Jiri A1 Domene, Xavier A1 Dvorsky, Miroslav A1 Ejtehadi, Hamid A1 Enrico, Lucas A1 Epikhin, Dmitrii A1 Eskelinen, Anu A1 Essl, Franz A1 Fan, Gaohua A1 Fantinato, Edy A1 Fazlioglu, Fatih A1 Fernández-Pascual, Eduardo A1 Ferrara, Arianna A1 Fidelis, Alessandra A1 Fischer, Markus A1 Flagmeier, Maren A1 Forte, T’ai G. W. A1 Fraser, Lauchlan H. A1 Fujinuma, Junichi A1 Furquim, Fernando F. A1 Garris, Berle A1 Garris, Heath W. A1 Giorgis, Melisa A. A1 Giusso del Galdo, Gianpietro A1 González-Robles, Ana A1 Good, Megan K. A1 Guardiola, Moisès A1 Guarino, Riccardo A1 Guerrero, Irene A1 Guillemot, Joannès A1 Güler, Behlül A1 Guo, Yinjie A1 Haesen, Stef A1 Hejda, Martin A1 Heleno, Ruben H. A1 Høye, Toke T. A1 Hrivnák, Richard A1 Huang, Yingxin A1 Hunter, John T. A1 Iakushenko, Dmytro A1 Ibáñez, Ricardo A1 Ingerpuu, Nele A1 Irl, Severin D. H. A1 Janíková, Eva A1 Jansen, Florian A1 Jeltsch, Florian A1 Jentsch, Anke A1 Jiménez-Alfaro, Borja A1 Jõks, Madli A1 Jouri, Mohammad H. A1 Karami, Sahar A1 Katal, Negin A1 Kelemen, András A1 Khairullin, Bulat I. A1 Khuroo, Anzar A. A1 Komatsu, Kimberly J. A1 Konečná, Marie A1 Kook, Ene A1 Korell, Lotte A1 Koroleva, Natalia A1 Korznikov, Kirill A. A1 Kozhevnikova, Maria V. A1 Kozub, Łukasz A1 Laanisto, Lauri A1 Lager, Helena A1 Lanta, Vojtech A1 Lasagno, Romina G. A1 Lembrechts, Jonas J. A1 Li, Liping A1 Lisner, Aleš A1 Liu, Houjia A1 Liu, Kun A1 Liu, Xuhe A1 Lucas-Borja, Manuel Esteban A1 Ludewig, Kristin A1 Lukács, Katalin A1 Luther-Mosebach, Jona A1 Macek, Petr A1 Marignani, Michela A1 Michalet, Richard A1 Miglécz, Tamás A1 Moeslund, Jesper Erenskjold A1 Moeys, Karlien A1 Montesinos, Daniel A1 Moreno-Jiménez, Eduardo A1 Moysiyenko, Ivan A1 Mucina, Ladislav A1 Muñoz-Rojas, Miriam A1 Murillo, Raytha A. A1 Nambahu, Sylvia M. A1 Neuenkamp, Lena A1 Normand, Signe A1 Nowak, Arkadiusz A1 Nuche, Paloma A1 Oja, Tatjana A1 Onipchenko, Vladimir G. A1 Pachedjieva, Kalina L. A1 Paganeli, Bruno A1 Peco, Begoña A1 Peralta, Ana M. L. A1 Pérez-Haase, Aaron A1 Peri, Pablo L. A1 Petraglia, Alessandro A1 Peyre, Gwendolyn A1 Plaza-Álvarez, Pedro Antonio A1 Plue, Jan A1 Prentice, Honor C. A1 Prokhorov, Vadim E. A1 Radujković, Dajana A1 Rahmanian, Soroor A1 Reitalu, Triin A1 Ristow, Michael A1 Robin, Agnès A. A1 Robles, Ana Belén A1 Rodríguez Ginart, Daniel A. A1 Román, Raúl A1 Roos, Ruben E. A1 Rosati, Leonardo A1 Sádlo, Jiří A1 Salimbayeva, Karina A1 Sánchez De Dios, Rut A1 Sanchir, Khaliun A1 Sattler, Cornelia A1 Scasta, John D. A1 Schmiedel, Ute A1 Schrader, Julian A1 Schultz, Nick L. A1 Sellan, Giacomo A1 Serra-Diaz, Josep M. A1 Silan, Giulia A1 Skálová, Hana A1 Skobel, Nadiia A1 Sonkoly, Judit A1 Štajerová, Kateřina A1 Svitková, Ivana A1 Świerszcz, Sebastian A1 Tanentzap, Andrew J. A1 Tanentzap, Fallon M. A1 Tarifa, Rubén A1 Tejero, Pablo A1 Tekeev, Dzhamal K. A1 Tholin, Michael A1 Thormodsæter, Ruben S. A1 Tian, Yichen A1 Tokaryuk, Alla A1 Tölgyesi, Csaba A1 Tomaselli, Marcello A1 Tordoni, Enrico A1 Török, Péter A1 Tóthmérész, Béla A1 Toussaint, Aurèle A1 Touzard, Blaise A1 Trindade, Diego P. F. A1 Tsakalos, James L. A1 Türkiş, Sevda A1 Valencia, Enrique A1 Valerio, Mercedes A1 Valkó, Orsolya A1 Van Meerbeek, Koenraad A1 Vandvik, Vigdis A1 Villellas, Jesus A1 Virtanen, Risto A1 Vítková, Michaela A1 Vojík, Martin A1 von Hessberg, Andreas A1 von Oppen, Jonathan A1 Wagner, Viktoria A1 Wan, Ji-Zhong A1 Wang, Chun-Jing A1 Wani, Sajad A. A1 Weiss, Lina A1 Wevill, Tricia A1 Xiao, Sa A1 Zárate Martínez, Oscar A1 Zobel, Martin AB Anthropogenic biodiversity decline threatens the functioning of ecosystems and the many benefits they provide to humanity1. As well as causing species losses in directly affected locations, human influence might also reduce biodiversity in relatively unmodified vegetation if far-reaching anthropogenic effects trigger local extinctions and hinder recolonization. Here we show that local plant diversity is globally negatively related to the level of anthropogenic activity in the surrounding region. Impoverishment of natural vegetation was evident only when we considered community completeness: the proportion of all suitable species in the region that are present at a site. To estimate community completeness, we compared the number of recorded species with the dark diversity—ecologically suitable species that are absent from a site but present in the surrounding region2. In the sampled regions with a minimal human footprint index, an average of 35% of suitable plant species were present locally, compared with less than 20% in highly affected regions. Besides having the potential to uncover overlooked threats to biodiversity, dark diversity also provides guidance for nature conservation. Species in the dark diversity remain regionally present, and their local populations might be restored through measures that improve connectivity between natural vegetation fragments and reduce threats to population persistence. PB Springer Nature SN 0028-0836 YR 2025 FD 2025-04-02 LK https://hdl.handle.net/20.500.14352/122590 UL https://hdl.handle.net/20.500.14352/122590 LA eng NO Pärtel, M., Tamme, R., Carmona, C.P. et al. Global impoverishment of natural vegetation revealed by dark diversity. Nature 641, 917–924 (2025). https://doi.org/10.1038/s41586-025-08814-5 NO AcknowledgementsWe thank the Estonian Research Council (PRG609, PRG1065, PSG293 and PRG2142) and the Estonian Ministry of Education and Research (Centre of Excellence AgroCropFuture, TK200) for funding the DarkDivNet steering committee, and we thank the many students and volunteers who helped with the fieldwork. Acknowledgements related to specific authors can be found in the Supplementary Notes:Additional acknowledgements• Jonathan A. Bennett: NSERC Discovery Grant (RGPIN-2018-04952)• Alessandro Chiarucci: National Recovery and Resilience Plan (NRRP), Project "National Biodiversity FutureCenter—NBFC" (CUP J33C22001190001)• Maarja Öpik: Estonian Research Council (1789 FUNFARM)• Vistorina Amputu: German Federal Ministry of Education and Research (BMBF; FKZ 01LC1821B)• Isabelle Aubin: K. Good, C. Zimmerman• Francisco M. Azcárate: Project REMEDINAL (TE-CM S2018/EMT-4338)• Zoltán Bátori: National Research Development and Innovation Office (FK142428), János Bolyai ResearchScholarship of the Hungarian Academy of Sciences (BO/00298/21/8), New National Excellence Program of theMinistry for Culture and Innovation from the source of the National Research, Development and Innovation Fund(ÚNKP-23-5-SZTE-697)• Natalie Beenaerts: Ann Cuypers, Anne Habets, Siebe Indestege• Idoia Biurrun: Basque Government (IT1487-22)• Bazartseren Boldgiv: Taylor Family-Asia Foundation Endowed Chair in Ecology and Conservation Biology• Francis Q. Brearley: Manchester Metropolitan University, Percy Sladen Memorial Fund• C. Guillermo Bueno: Ramon y Cajal Fellowship (RYC2021-032533-I), consolidator grant "Mycolegium" from theSpanish Ministry of Science, Innovation and University (CNS2023-143989)• James F. Cahill: NSERC Discovery Grant• Juan A. Campos: Basque Government (IT1487-22)• Christopher Carcaillet: Olivier Blarquez• Jan Čuda: Czech Academy of Sciences, long-term research development project (RVO 67985939)• Mariana Dairel: FAPESP (2019/09659-1)• John Davison: Estonian Research Council (PRG2584)• Balázs Deák: Hungarian National Research, Development and Innovation Office (NKFIH FK 135329, NKFIH KKP144096)• Jiri Dolezal: Czech Science Foundation (24-11954S), MSMT INTER-ACTION (LUAUS24258)• Anu Eskelinen: Research Council of Finland (project no 351089)• Franz Essl: Austrian Science Foundation FWF (I 5825-B)• Fatih Fazlioglu: Ordu University Research Council (A-2028)• Arianna Ferrara: Ente Parco Nazionale Foreste Casentinesi (PhD project "Conservation of natural vegetationdynamics within the National Park of Casentinesi Forests")• Alessandra Fidelis: São Paulo Research Foundation (FAPESP 2015/06743-0), National Council for Scientific andTechnological Development (CNPq 312689/2021-7), permit COTEC nº 480/2018 D67/2018 PH (SMA/SP), HeloizaZirondi, Vagner Zanzarini, Cassy Anne Rodrigues• Markus Fischer: German Research Foundation (DFG Priority Programme 1374, "Biodiversity Exploratories")• Lauchlan H. Fraser: Natural Sciences and Engineering Research Council of Canada Industrial Research Chair inEcosystem Reclamation• Fernando F. Furquim: Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES), Brasil (Financecode 001)• Heath W. Garris: Covenant College• Ana González-Robles: Junta de Andalucía (Delegación Territorial de Agricultura, Ganadería, Pesca y DesarrolloSostenible en Almería), Parque Natural Cabo de Gata-Níjar and 'Playas y Cortijos family' for facilitating permits todo the field samplings, J.L. González Rebollar• Stef Haesen: FLOF fellowship (3E190655) of the KU Leuven• Martin Hejda: Czech Academy of Sciences, long-term research development project (RVO 67985939)• Ruben H. Heleno: Portuguese Foundation for Science and Technology (UIDB/04004/2020,10.54499/LA/P/0092/2020)• Toke T. Høye: Independent Research Fund Denmark (8021-00423B)• Yingxin Huang: Strategic Priority Research Program of the Chinese Academy of Sciences (XDA28110202), projectsupported by the National Natural Science Foundation of China (42471071 ), China Jilin Provincial Major Scienceand Technology Project (20230303008SF)• John T. Hunter: Vanessa Hunter• Ricardo Ibáñez: University of Navarra (project “Red de Observatorios de la Biodiversidad de Navarra (ROBIN)”)• Borja Jiménez-Alfaro: Spanish Research Agency (PID2019-108636GA-100)• András Kelemen: János Bolyai Research Scholarship of the Hungarian Academy of Sciences (BO/00713/19)12• Kimberly J. Komatsu: Smithsonian Institution• Natalia Koroleva: Ekaterina Kopeina• Lauri Laanisto: Estonian Academy of Sciences (research professorship for Arctic studies), Adam MickiewiczUniversity Polar Station (Petuniabukta)• Vojtech Lanta: Czech Science Foundation (23-07533S)• Jonas J. Lembrechts: Research Foundation Flanders (G018919N, 12P1819N), BiodivERsA+ (The ASICS Project)• Liping Li: National Natural Science Foundation of China (82274041)• Kristin Ludewig: Harry Stöckmann, Jutta Krüger• Petr Macek: Czech Science Foundation (GA17-20839S)• Jesper Erenskjold Moeslund: Aage V. Jensen Naturfond ("Dark diversity in nature management")• Eduardo Moreno-Jiménez: Spanish Ministry of Science and Innovation (PID2020-116578RB-I00)• Ladislav Mucina: Iluka Chair at the Murdoch University, Perth, Australia• Miriam Muñoz-Rojas: Ministry of Science and Innovation, Spain (RYC2020-029255-I, TED2021-132332A-C22,PID2021-123097OA-I00)• Lena Neuenkamp: German Science Foundation (DFG Priority Programme 1374 "Biodiversity Exploratories")• Signe Normand: Independent Research Fund Denmark (7027-00133B ), Novo Nordisk Foundation ChallengeProgram, Matilde Mikkeline Holst Stoltze• Arkadiusz Nowak: National Science Centre, Poland (2019/35/B/NZ8/03358)• Begoña Peco: Helios Sainz• Alessandro Petraglia: COMP-HUB and COMP-R Initiatives, funded by the ‘Departments of Excellence’ program ofthe Italian Ministry for University and Research (MIUR, 2018-2022 and MUR, 2023-2027)• Pedro Antonio Plaza Álvarez: Regional Government of Castilla-La Mancha and the European Social Fund Plus(ESF+)• Jan Plue: Future Research Leader grant by FORMAS (2018-00961)• Triin Reitalu: Estonian Research Council (PRG874), Estonian Ministry of Education and Research, Centre ofExcellence for Sustainable Land Use (TK232)• Michael Ristow: Merlin Schäfer, Marieke Bittner• Ana Belén Robles: Project Research on Forage Flora of Baetica (Ref. 201640E002) Spanish Council for ScientificResearch (CSIC) Agency• Raúl Román: European Union Horizon 2020 research and innovation programme under the Marie SkłodowskaCurie IF-GF Actions (MICROBIOCLIM, GA nr. 101028323)• Jiří Sádlo: Technology Agency of the Czech Republic, project DivLand (TACR SS02030018), Czech Academy ofSciences, long-term research development project (RVO 67985939)• Khaliun Sanchir: Taylor Family-Asia Foundation Endowed Chair in Ecology and Conservation Biology• Ute Schmiedel: German Federal Ministry of Education and Research (01LC0024, 01LC0024A, 01LC0624A2,01LG1201A, 01LG1201N), the Soebatsfontein community (that gave permission to work on their farmland), JonaLuther-Mosebach, Inga U. Röwer, Reginald Christiaan• Josep M. Serra-Diaz: Agence Nationale de la Recherche programme Jeune Chercheur (ANR-21-CE32-0003), RemyJoly, Anna Schmidt• Hana Skálová: Technology Agency of the Czech Republic, project DivLand (TACR SS02030018), Technology Agencyof the Czech Republic, BiodivClim Call 2019, project ASICS (TACR SS70010001), Czech Academy of Sciences, longterm research development project (RVO 67985939)• Judit Sonkoly: National Research, Development and Innovation Office (PD 137747), Bolyai János Scholarship ofthe Hungarian Academy of Sciences (BO/00587/23/8)• Kateřina Štajerová: Czech Academy of Sciences, long-term research development project (RVO 67985939), AnnaLučanová• Ivana Svitková: Scientific Grant Agency of the Ministry of Education, Science, Research and Sport of the SlovakRepublic and Slovak Academy of Sciences (VEGA 2/0031/17)• Andrew J. Tanentzap: Canada Research Chairs Program and Natural Sciences and Engineering Research Council ofCanada (NSERC; RGPIN-2023-03977)• Pablo Tejero: JACA Herbarium (IPE-CSIC)• Csaba Tölgyesi: National Research Development and Innovation Office (K146137)• Enrico Tordoni: Estonian Research Council (PSG1021)• Péter Török: National Research Found of Hungary (KKP 144068)• Diego P. F. Trindade: Estonian Research Council, Estonian Research Council (PUTJD1228)• Enrique Valencia: Community of Madrid in the framework of the Multiannual Agreement with the Rey JuanCarlos University in line of action 1, “Encouragement of Young Phd students investigation” (M2165, INTRANESTI)• Mercedes Valerio: Departamento de Educación, Gobierno de Navarra (Ayudas predoctorales para la realizaciónde programas de doctorado de interés para Navarra; Plan de Formación y de I + D 2018)• Orsolya Valkó: Hungarian National Research, Development and Innovation Office (KKP 144096)13• Michaela Vítková: Technology Agency of the Czech Republic, project DivLand (TACR SS02030018), TechnologyAgency of the Czech Republic, BiodivClim Call 2019, project ASICS (TACR SS70010001), Czech Academy ofSciences, long-term research development project (RVO 67985939)• Martin Vojík: Czech Academy of Sciences, long-term research development project (RVO 67985939), TechnologyAgency of the Czech Republic, project DivLand (TACR SS02030018)• Jonathan von Oppen: The Independent Research Fund of Denmark (7027-00133B)• Viktoria Wagner: Discovery Grant from the Canadian Natural Science and Engineering Research Council(RES0043426), Competitive Grant from the University of Alberta Rangeland Research Institute (RES0045021),Startup Grant from the University of Alberta• Ji-Zhong Wan: Hundred Talent Program of Chinese Academy of Sciences (No. E4K2150150)• Sajad A. 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