[1] Bangor Univ, Sch Ocean Sci, Menai Bridge LL59 5AB, Anglesey, Wales
[2] Royal Netherlands Inst Sea Res, NL-1790 AB Den Burg, Texel, Netherlands
[3] Free Univ Amsterdam, NL-1081 HV Amsterdam, Netherlands
[4] Univ N Carolina, Ctr Marine Sci, Wilmington, NC 28409 USA
[5] Univ N Carolina, Dept Marine Sci, Chapel Hill, NC 27599 USA