Life-history theory predicts that current behaviour affects future reproduction, implying that animals should optimise their escape strategies to reflect fitness costs and benefits of premature escape. Both costs and benefits of escape may change temporally with important consequences for the evolution of escape strategies. Moreover, escape strategies of species may differ according to their positions on slow-fast pace of life gradients. We studied risk-taking in long-distance migratory animals, waders (Charadriiformes), during the annual cycle, i.e., breeding in Europe, stopover in the Middle East and wintering in tropical Africa. Phylogenetically informed comparative analyses revealed that risk-taking (measured as flight initiation distance, FID) changed significantly over the year, being lowest during breeding and peaking at stopover sites. Similarly, relationships between risk-taking and life-history traits changed among stages of the annual cycle. While risk-taking significantly decreased with increasing body mass during breeding, risk-taking-body mass relationship became marginally significant in winter and disappeared during migration. The positive trend of risk-taking along slow-fast pace of life gradient measured as adult survival was only found during breeding. The season-dependent relationships between risk-taking and life history traits suggest that migrating animals respond to fluctuating environments by adopting behavioural plasticity.

Arctic Centre University of Lapland PO Box 122 96101 Rovaniemi Finland

Ben Gurion University of the Negev Eilat Campus P O Box 272 Eilat 88000 Israel

Department of Biogeography and Global Change Museo Nacional de Ciencias Naturales CSIC c Serrano 115bis E 28006 Madrid Spain

Department of Zoology Faculty of Science Charles University Viničná 7 128 43 Praha 2 Czech Republic

Ecologie Systématique Evolution Université Paris Sud CNRS AgroParisTech Université Paris Saclay F 91405 Orsay Cedex France

Faculty of Biological Sciences University of Zielona Góra Prof Z Szafrana 1 65 516 Zielona Góra Poland

Institute of Vertebrate Biology Czech Academy of Sciences Květná 8 603 65 Brno Czech Republic

Institute of Zoology Poznań University of Life Sciences Wojska Polskiego 71C 60 625 Poznań Poland

Laboratory and Museum of Evolutionary Ecology Department of Ecology Faculty of Humanities and Natural Sciences University of Presov 17 novembra 1 080 01 Prešov Slovakia

Rabin High School Yotam Street 51 Eilat 8820301 Israel

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