[1] NOAA, Natl Marine Mammal Lab, Alaska Fisheries Sci Ctr, Natl Marine Fisheries Serv, Seattle, WA 98115 USA
[2] Univ St Andrews, Sea Mammal Res Unit, St Andrews KY16 8LB, Fife, Scotland
[3] Univ St Andrews, Ctr Res Ecol & Environm Modelling, St Andrews KY16 9LZ, Fife, Scotland
[4] Univ St Andrews, Sch Math & Stat, St Andrews KY16 9LZ, Fife, Scotland
[5] Univ Glasgow, Inst Biodivers Anim Hlth & Comparat Med, Glasgow G12 8QQ, Lanark, Scotland