1. School of Mathematics and Statistics, University of Glasgow, Glasgow G12 8SQ, UK
2. Institute of Biodiversity, Animal Health and Comparative Medicine, University of Glasgow, Glasgow G12 8QQ, UK
3. Department of Collective Behaviour, Max Planck Institute for Ornithology, 78464 Konstanz, Germany
4. Chair of Biodiversity and Collective Behaviour, Department of Biology, University of Konstanz, 78464 Konstanz, Germany
5. Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ 08544, USA