[1] Univ Tasmania, Inst Marine & Antarctic Studies, Hobart, Tas 7000, Australia
[2] CSIRO, Castray Esplanade, Hobart, Tas, Australia
[3] Univ Tasmania, Inst Marine & Antarctic Studies, Hobart, Tas, Australia
[4] Univ St Andrews, Sch Biol, Scottish Oceans Inst, St Andrews KY16 8LB, Fife, Scotland