1. Department of Evolutionary Biology, Ecology and Environmental Sciences, University of Barcelona, 08028 Barcelona, Spain
2. Department of Zoology, University of Oxford, Oxford OX1 3SZ Oxford, United Kingdom
3. Evolutionary Biodemography Laboratory, Max Planck Institute for Demographic Research, Rostock 18057, Germany
4. Centre for Biodiversity and Conservation Science, University of Queensland, St Lucia, Queensland 4072, Australia