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Contrasting thermophilization among forests, grasslands and alpine summits

Yue, Kai ORCID: https://orcid.org/0000-0002-7709-8523; Vangansbeke, Pieter ORCID: https://orcid.org/0000-0002-6356-2858; Myers-Smith, Isla H. ORCID: https://orcid.org/0000-0002-8417-6112; Waller, Donald M. ORCID: https://orcid.org/0000-0001-5377-3929; Verheyen, Kris ORCID: https://orcid.org/0000-0002-2067-9108; Bernhardt-Römermann, Markus ORCID: https://orcid.org/0000-0002-2740-2304; Baeten, Lander ORCID: https://orcid.org/0000-0003-4262-9221; Staude, Ingmar R. ORCID: https://orcid.org/0000-0003-2306-8780; Bjorkman, Anne D.; Hédl, Radim ORCID: https://orcid.org/0000-0002-6040-8126; Andrews, Christopher ORCID: https://orcid.org/0000-0003-2428-272X; Barni, Elena ORCID: https://orcid.org/0000-0001-7256-0064; Becker, Thomas; Becker-Scarpitta, Antoine ORCID: https://orcid.org/0000-0001-9241-091X; Benito-Alonso, José-Luis ORCID: https://orcid.org/0000-0003-1086-8834; Bennie, Jonathan ORCID: https://orcid.org/0000-0003-4394-2041; Berki, Imre; Blüml, Volker ORCID: https://orcid.org/0000-0002-9910-3469; Brunet, Jörg ORCID: https://orcid.org/0000-0003-2667-4575; Bullock, James M. ORCID: https://orcid.org/0000-0003-0529-4020; Van Calster, Hans ORCID: https://orcid.org/0000-0001-8595-8426; Carbognani, Michele ORCID: https://orcid.org/0000-0001-7701-9859; Chudomelová, Markéta; Closset-Kopp, Déborah; Dan Turtureanu, Pavel ORCID: https://orcid.org/0000-0002-7422-3106; Daskalova, Gergana N. ORCID: https://orcid.org/0000-0002-5674-5322; Decocq, Guillaume; Dick, Jan ORCID: https://orcid.org/0000-0002-4180-9338; Diekmann, Martin; Dirnböck, Thomas ORCID: https://orcid.org/0000-0002-8294-0690; Durak, Tomasz ORCID: https://orcid.org/0000-0003-4053-3699; Eriksson, Ove; Erschbamer, Brigitta; Graae, Bente Jessen; Heinken, Thilo ORCID: https://orcid.org/0000-0002-1681-5971; Hermy, Martin ORCID: https://orcid.org/0000-0002-5403-0139; Horchler, Peter; Jandt, Ute ORCID: https://orcid.org/0000-0002-3177-3669; Jaroszewicz, Bogdan ORCID: https://orcid.org/0000-0002-2042-8245; Kanka, Róbert; Kollár, Jozef; Kopecký, Martin ORCID: https://orcid.org/0000-0002-1018-9316; Kudernatsch, Thomas; Lamprecht, Andrea ORCID: https://orcid.org/0000-0002-8719-026X; Lenoir, Jonathan ORCID: https://orcid.org/0000-0003-0638-9582; Macek, Martin ORCID: https://orcid.org/0000-0002-5609-5921; Malicki, Marek; Máliš, František ORCID: https://orcid.org/0000-0003-2760-6988; Michelsen, Ottar ORCID: https://orcid.org/0000-0001-7141-1948; Mitchell, Fraser; Naaf, Tobias ORCID: https://orcid.org/0000-0002-4809-3694; Nagel, Thomas A. ORCID: https://orcid.org/0000-0002-4207-9218; Newman, Miles; Newton, Adrian C.; Nicklas, Lena; Oddi, Ludovica; Orczewska, Anna; Orsenigo, Simone ORCID: https://orcid.org/0000-0003-0348-9115; Ortmann-Ajkai, Adrienne; Ouden, Jan den; Pauli, Harald; Peterken, George; Petřík, Petr ORCID: https://orcid.org/0000-0001-8518-6737; Pielech, Remigiusz ORCID: https://orcid.org/0000-0001-8879-3305; Puşcaş, Mihai; Randin, Christophe; Reczyńska, Kamila ORCID: https://orcid.org/0000-0002-0938-8430; Rixen, Christian ORCID: https://orcid.org/0000-0002-2486-9988; Schei, Fride Høistad ORCID: https://orcid.org/0000-0001-8083-4045; Schmidt, Wolfgang ORCID: https://orcid.org/0000-0001-5356-4684; Šebesta, Jan; Stachurska-Swakon, Alina ORCID: https://orcid.org/0000-0003-0381-4520; Standovár, Tibor ORCID: https://orcid.org/0000-0002-4686-3456; Świerkosz, Krzysztof ORCID: https://orcid.org/0000-0002-5145-178X; Teleki, Balázs; Theurillat, Jean-Paul ORCID: https://orcid.org/0000-0002-1843-5809; Ursu, Tudor-Mihai ORCID: https://orcid.org/0000-0002-4898-6345; Vanneste, Thomas; Vellend, Mark; Vergeer, Philippine; Vild, Ondřej; Villar, Luis; Vittoz, Pascal ORCID: https://orcid.org/0000-0003-4218-4517; Winkler, Manuela ORCID: https://orcid.org/0000-0002-8655-9555; Wipf, Sonja ORCID: https://orcid.org/0000-0002-3492-1399; Wu, Fuzhong ORCID: https://orcid.org/0000-0002-8164-6078; Zhang, Shengmin ORCID: https://orcid.org/0000-0001-8770-6935; De Frenne, Pieter ORCID: https://orcid.org/0000-0002-8613-0943. 2026 Contrasting thermophilization among forests, grasslands and alpine summits. Nature. 10.1038/s41586-025-09622-7

Abstract
Climate warming is shifting biological communities, with warmth-demanding species being favoured at the expense of cold-adapted species in a process referred to as thermophilization. Because biodiversity responses often lag behind climate warming, climatic debts are accumulating in many ecosystems across the world. Although we might expect that thermophilization and climatic debts will vary among habitats, standardized quantification across ecosystems is lacking. Here we analysed multidecadal data from 6,067 resurveyed vegetation plots over 12–78 years in forests, grasslands and on alpine summits across Europe. We demonstrate that forest understory and grassland plant communities experienced positive thermophilization, although not significantly different from zero. By contrast, alpine summit vegetation showed much stronger (up to five times) and significant thermophilization. Thermophilization was driven largely by increases in warmth-demanding species in grasslands, by declines in cold-adapted species on alpine summits and by both processes in forests. Significant climatic debts have accumulated in forests and alpine summits, but less so in grasslands, with debts positively correlated with macroclimate temperature changes. Our findings uncover divergent thermophilization trajectories and increasing climatic debts across ecosystems. Moreover, we highlight the mechanisms that enable some communities to track climate change more closely than others and provide a basis for projecting future shifts in plant communities under accelerating climate warming.
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