1. Scottish Oceans Institute, School of Biology, University of St Andrews, St Andrews, Fife KY16 8LB, UK
2. National Marine Fisheries Service, 28 Tarzwell Drive, Narragansett, RI 02882, USA
3. Freshwater Laboratory Field Station, Inchbraoch House, South Quay, Ferryden, Montrose, Angus DD10 9SL, UK
4. Cefas, Pakefield Road, Lowestoft, Suffolk NR33 0HT, UK
5. Centre for Evolution, Genes and Genomics, School of Biology, University of St Andrews, St Andrews, Fife KY16 9TH, UK