[1] Univ Auckland, Inst Marine Sci, Auckland 1142, New Zealand
[2] Natl Inst Water & Atmospher Res, Hamilton 3251, New Zealand
[3] Albert Ludwig Univ Freiburg, Freiburg, Germany
[4] Nord Univ, Fac Biosci & Aquaculture, Postboks 1490, N-8049 Bodo, Norway
[5] Univ Auckland, Sch Environm, Auckland, New Zealand