The extent of vegetation openness in past European landscapes is widely debated. In particular, the temperate forest biome has traditionally been defined as dense, closed-canopy forest; however, some argue that large herbivores maintained greater openness or even wood-pasture conditions. Here, we address this question for the Last Interglacial period (129,000-116,000 years ago), before Homo sapiens-linked megafauna declines and anthropogenic landscape transformation. We applied the vegetation reconstruction method REVEALS to 96 Last Interglacial pollen records. We found that light woodland and open vegetation represented, on average, more than 50% cover during this period. The degree of openness was highly variable and only partially linked to climatic factors, indicating the importance of natural disturbance regimes. Our results show that the temperate forest biome was historically heterogeneous rather than uniformly dense, which is consistent with the dependency of much of contemporary European biodiversity on open vegetation and light woodland.

Center for Ecological Dynamics in a Novel Biosphere Department of Biology Aarhus University Ny Munkegade 114 DK 8000 Aarhus C Denmark

Center for Landscape Research in Sustainable Agricultural Futures Department of Biology Aarhus University Ny Munkegade 114 DK 8000 Aarhus C Denmark

Center for Sustainable Landscapes under Global Change Department of Biology Aarhus University Ny Munkegade 114 DK 8000 Aarhus C Denmark

CEREGE CNRS IRD Europôle de l'Arbois BP 80 F 13545 Aix en Provence France

Climate Change Ecology Research Unit Faculty of Geographical and Geological Sciences Adam Mickiewicz University Poznań Bogumiła Krygowskiego 10 Poznań 61 680 Poland

Department of Botany Charles University Prague Czechia

Department of Environmental Geography CNRS UMR GEODE 5602 University Toulouse Jean Jaurès Toulouse France

Department of Geography and Environment School of Geosciences University of Aberdeen UK

Department of Geography Royal Holloway University of London Egham UK

Department of Geography University of Cambridge Cambridge CB2 3EN UK

Department of Geology and Geomorphology University of Łódź Narutowicza 88 90 139 Łódź Poland

Department of Geology Lund University Sölvegatan 12 SE 223 62 Lund Sweden

Department of Geosciences and Geography University of Helsinki Helsinki Finland

Department of Physical Geography and Quaternary Geology Stockholm University SE 106 91 Stockholm Sweden

Dipartimento di Biologia Ambientale University of Rome 'La Sapienza' Rome Italy

Faculty of Archaeology Leiden University Einsteinweg 2 2333 CC Leiden Netherlands

Faculty of Earth Sciences and Spatial Management Nicolaus Copernicus University in Toruń Lwowska 1 87 100 Toruń Poland

Faculty of Geology University of Warsaw Warsaw Poland

Faculty of Natural Sciences University of Silesia Będzińska 60 41 200 Sosnowiec Poland

Faculty of Science Department of Earth Sciences Vrije Universiteit Amsterdam Amsterdam Netherlands

Gymnasieskolan Knut Hahn Blasius Königsgatan 27 37232 Ronneby Sweden

Independent researcher Soloviny str 4 1 224 117593 Moscow Russia

Institute for Environmental Studies Vrije Universiteit Amsterdam Amsterdam Netherlands

Institute of Plant Sciences and Oechger Centre for Climate Change Research University of Bern Altenbergrain 21 3013 Bern Switzerland

Instituto Pirenaico de Ecología IPE CSIC Avda Montañana 1005 50059 Zaragoza Spain

Laboratory of Paleobotany Department of Stratigraphical Geology Institute of Geological Sciences University of Wroclaw Cybulskiego 34 50 205 Wroclaw Poland

Landesamt für Bergbau Geologie und Rohstoffe Inselstraße 26 03046 Cottbus Germany

Leuphana University Lüneburg Institute of Ecology Lüneburg Germany

Maria Curie Sklodowska University Institute of Earth and Environmental Sciences Al Krasnicka 2 d 20 718 Lublin Poland

Mediterranean Institute of Marine and Terrestrial Biodiversity and Ecology Aix Marseille University Marseille France

Nature Research Centre Institute of Geology and Geography Akademijos 2 LT 08412 Vilnius Lithuania

Ontario Geological Survey 933 Ramsey Lake Road Sudbury ON P3E 6B5 Canada

Polish Geological Institute 00 975 Warsaw Poland

Polish Geological Institute National Research Institute Carpathian Branch Skrzatów 1 31 560 Kraków Poland

Polish Geological Institute National Research Institute Marine Geology Branch ul Kościerska 5 80 328 Gdańsk Poland

School of Geography Earth and Environmental Sciences University of Plymouth Plymouth UK

Section for Ecoinformatics and Biodiversity Department of Biology Aarhus University Ny Munkegade 114 DK 8000 Aarhus C Denmark

TNO Geological Survey of the Netherlands Postbus 80015 3508 TA Utrecht Netherlands

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