Centre for Evolution, Genes and Genomics, School of Biology, University of St Andrews, St Andrews, Fife, KY16 9TH, UK.|Centre for Evolution, Genes and Genomics, School of Biology, University of St Andrews, St Andrews, Fife, KY16 9TH, UK.|Centre for Evolution, Genes and Genomics, School of Biology, University of St Andrews, St Andrews, Fife, KY16 9TH, UK.|Centre for Evolution, Genes and Genomics, School of Biology, University of St Andrews, St Andrews, Fife, KY16 9TH, UK.|Centre for Evolution, Genes and Genomics, School of Biology, University of St Andrews, St Andrews, Fife, KY16 9TH, UK.