1. Institute of Biological and Environmental Sciences, School of Biological Sciences, University of Aberdeen, Aberdeen AB24 2TZ, UK
2. Glasgow Ageing Research Network (GARNER), Institute of Biodiversity, Animal Health and Comparative Medicine, University of Glasgow, Glasgow AB24 2TZ, UK
3. College of Life and Environmental Sciences, University of Exeter, Penryn Campus, TR10 9EZ, UK
4. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh EH9 3JG, UK