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1) Centre for Social Evolution, Department of Biology, University of Copenhagen, Copenhagen, 14 Denmark 15 2) Present address: Centre of Excellence in Biological interactions, Department of Biosciences, 16 University of Helsinki, Finland 17 3) Laboratoire d'Ethologie Expérimentale et Comparée (LEEC), Université Paris 13, 18 Villetaneuse, France 19 4) Institut de Recherche sur la Biologie de l'Insecte (IRBI), UMR CNRS 6035, Université 20 François-Rabelais, Tours, France 21 http://jeb.biologists.org/lookup/doi/10.1242/jeb.087007 Access the most recent version at J Exp Biol Advance Online Articles. First posted online on 16 May 2013 as doi:10.1242/jeb.087007