Much research on large herbivore movement has focused on the annual scale to distinguish between resident and migratory tactics, commonly assuming that individuals are sedentary at the within-season scale. However, apparently sedentary animals may occupy a number of sub-seasonal functional home ranges (sfHR), particularly when the environment is spatially heterogeneous and/or temporally unpredictable. The roe deer (Capreolus capreolus) experiences sharply contrasting environmental conditions due to its widespread distribution, but appears markedly sedentary over much of its range. Using GPS monitoring from 15 populations across Europe, we evaluated the propensity of this large herbivore to be truly sedentary at the seasonal scale in relation to variation in environmental conditions. We studied movement using net square displacement to identify the possible use of sfHR. We expected that roe deer should be less sedentary within seasons in heterogeneous and unpredictable environments, while migratory individuals should be seasonally more sedentary than residents. Our analyses revealed that, across the 15 populations, all individuals adopted a multi-range tactic, occupying between two and nine sfHR during a given season. In addition, we showed that (i) the number of sfHR was only marginally influenced by variation in resource distribution, but decreased with increasing sfHR size; and (ii) the distance between sfHR increased with increasing heterogeneity and predictability in resource distribution, as well as with increasing sfHR size. We suggest that the multi-range tactic is likely widespread among large herbivores, allowing animals to track spatio-temporal variation in resource distribution and, thereby, to cope with changes in their local environment.

Bavarian State Institute of Forestry Hans Carl von Carlowitz Platz 1 85354 Freising Germany

Biodiversity and Molecular Ecology Department Research and Innovation Centre Fondazione Edmund Mach Via Mach 1 38010 San Michele all'Adige TN Italy

CEFS Université de Toulouse INRA Castanet Tolosan France

Centre for Ecological and Evolutionary Synthesis Department of Biosciences University of Oslo Blindern P O Box 1066 NO 0316 Oslo Norway

Chair of Wildlife Ecology and Management University of Freiburg Tennenbacher Straße 4 79106 Freiburg Germany Forest Research Institute of Baden Wuerttemberg Wildlife Ecology Wonnhaldestraße 4 79100 Freiburg Germany

CNRS Univ Montpellier EPHE IRD Univ Paul Valéry Montpellier 3 34293 Montpellier France

Department of Biodiversity Research Global Change Research Institute CAS Academy of Sciences of the Czech Republic Brno Czech Republic Šumava National Park Vimperk Czech Republic

Department of Conservation and Research Bavarian Forest National Park Freyunger Str 2 94481 Grafenau Germany Chair of Wildlife Ecology and Management University of Freiburg Tennenbacher Straße 4 79106 Freiburg Germany

Department of Evolutionary Biology and Environmental Studies University of Zurich Winterthurerstrasse 190 8057 Zurich Germany

Department of Integrative Biology and Biodiversity Research Institute of Wildlife Biology and Game Management University of Natural Resources and Life Sciences Vienna Austria

Freelance Consultant Milan Italy

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

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Office National de la Chasse et de la Faune Sauvage Direction Recherche et Expertise Unité Ongulés Sauvages Monfort 01330 Birieux France

Research Ecology and Environment Dimension Via Enrico Bindi 14 51100 Pistoia PT Italy

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