[1] Univ Edinburgh, Sch Biol Sci, Inst Evolutionary Biol, Edinburgh EH9 3JT, Midlothian, Scotland
[2] Australian Inst Marine Sci, Darwin, NT 8010, Australia
[3] Univ Auckland, Sch Biol Sci, Leigh Marine Lab, Auckland 1, New Zealand
[4] Curtin Univ Technol, Ctr Marine Sci & Technol, Perth, WA 6485, Australia