1. Primary objectives of national parks usually include both, the protection of natural processes and species conservation. When these objectives conflict, as occurs because of the cascading effects of large mammals (i.e., ungulates and large carnivores) on lower trophic levels, park managers have to decide upon the appropriate management while considering various local circumstances. 2. To analyse if ungulate management strategies are in accordance with the objectives defined for protected areas, we assessed the current status of ungulate management across European national parks using the naturalness concept and identified the variables that influence the management. 3. We collected data on ungulate management from 209 European national parks in 29 countries by means of a large-scale questionnaire survey. Ungulate management in the parks was compared by creating two naturalness scores. The first score reflects ungulate and large carnivore species compositions, and the second evaluates human intervention on ungulate populations. We then tested whether the two naturalness score categories are influenced by the management objectives, park size, years since establishment, percentage of government-owned land, and human impact on the environment (human influence index) using two generalized additive mixed models. 4. In 67.9% of the national parks, wildlife is regulated by culling (40.2%) or hunting (10.5%) or both (17.2%). Artificial feeding occurred in 81.3% of the national parks and only 28.5% of the national parks had a non-intervention zone covering at least 75% of the area. Furthermore, ungulate management differed greatly among the different countries, likely because of differences in hunting traditions and cultural and political backgrounds. Ungulate management was also influenced by park size, human impact on the landscape, and national park objectives, but after removing these variables from the full model the reduced models only showed a small change in the deviance explained. In areas with higher anthropogenic pressure, wildlife diversity tended to be lower and a higher number of domesticated species tended to be present. Human intervention (culling and artificial feeding) was lower in smaller national parks and when park objectives followed those set by the International Union for the Conservation of Nature (IUCN). 5. Our study shows that many European national parks do not fulfil the aims of protected area management as set by IUCN guidelines. In contrast to the USA and Canada, Europe currently has no common ungulate management policy within national parks. This lack of a common policy together with differences in species composition, hunting traditions, and cultural or political context has led to differences in ungulate management among European countries. To fulfil the aims and objectives of national parks and to develop ungulate management strategies further, we highlight the importance of creating a more integrated European ungulate management policy to meet the aims of national parks.

ARCTUROS Civil Society for the Protection and Management of Wildlife and the Natural Environment Aetos 53075 Florina Greece

Biotechnical Faculty University of Ljubljana Jamnikarjeva 101 1000 Ljubljana Slovenia

Cairngorms National Park Authority 14 The Square Grantown on Spey PH26 3HG UK

Chair of Wildlife Ecology and Management Albert Ludwigs University Freiburg Tennenbacher Straβe 4 79106 Freiburg Germany

Chair of Zoology 3 University of Würzburg Am Hubland 97074 Würzburg Germany

Department of Biodiversity and Molecular Ecology Fondazione Edmund Mach Via Edmund Mach 1 38010 San Michele all'Adige Italy

Department of Biology and CESAM University of Aveiro Campus Universitário de Santiago 3810 193 Aveiro Portugal

Department of Bioscience Aarhus University Kalø Grenåvej 14 8410 Rønde Denmark

Department of Conservation and Research Bavarian Forest National Park Freyunger Straβe 2 94481 Grafenau Germany

Department of Environmental Sciences Resource Ecology Group Wageningen University Droevendaalsesteeg 3a 6708 PB Wageningen the Netherlands

Department of Fisheries Beekeeping Game Management and Special Zoology Faculty of Agriculture University of Zagreb Svetošimunska cesta 25 10000 Zagreb Croatia

Department of Forestry Hochschule Weihenstephan Triesdorf Hans Carl von Carlowitz Platz 3 85354 Freising Germany

Department of Geography Faculty of Mathematics and Natural Sciences Humboldt University Unter den Linden 6 10099 Berlin Germany

Department of Organismic and Evolutionary Biology Harvard University 26 Oxford Street Cambridge MA 02138 USA

Department of Primatology Max Planck Institute for Evolutionary Anthropology Deutscher Platz 6 04103 Leipzig Germany

Department of Research and Nature Protection Šumava National Park and PLA Administration Sušická 399 34192 Kašperské Hory Czech Republic

Department of Silviculture Faculty of Silviculture and Forest Engineering Transilvania University 1 Beethoven Lane 500123 Brașov Romania

Department of Veterinary Medicine University of Sassari Via Vienna 2 07100 Sassari Italy

Department of Visitor Management and National Park Monitoring Bavarian Forest National Park Freyunger Straβe 2 94481 Grafenau Germany

Department of Zoology and Anthropology Faculty of Biology Sofia University St Kliment Ohridski bul Tsar Osvoboditel 15 1164 Sofia Bulgaria

Faculty of Environmental Sciences and Natural Resource Management Norwegian University of Life Sciences Universiteitstunet 3 1433 Ås Norway

Faculty of Forestry and Wood Sciences Czech University of Life Sciences Prague Kamýcká 1176 16521 Prague Czech Republic

Forest Research Institute Bulevardul Eroilor Number 128 Voluntari Ilfov 077190 Romania

German Centre for Integrative Biodiversity Research Deutscher Platz 5e 04103 Leipzig Germany

Grimsö Wildlife Research Station Department of Ecology Swedish University of Agricultural Sciences 73091 Riddarhyttan Sweden

Latvian State Forest Research Institute SILAVA Rīgas iela 111 2169 Salaspils Latvia

Mammal Research Institute Polish Academy of Sciences Stoczek 1 17 230 Białowieża Poland

Metsähallitus Parks and Wildlife Finland Lars Sonckin kaari 14 02600 Espoo Finland

Nature Research Centre Akademijos 2 08412 Vilnius Lithuania

Norwegian Institute for Nature Research PO Box 5685 Torgard 7485 Trondheim Norway

Swiss National Park Chastè Planta Wildenberg 7530 Zernez Switzerland

The Environmental Board Narva maantee 7a 15172 Tallinn Estonia

Ungulate Research Unit Cátedra de Recursos Cinegéticos y Piscícolas Universidad de Córdoba 14071 Córdoba Spain

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