1. Department of Marine Biogeochemistry and Toxicology, Netherlands Institute for Sea Research, P.O. Box 59, 1790 AB Den Burg, The Netherlands, Department of Marine Ecology, Netherlands Institute for Sea Research, P.O. Box 59, 1790 AB Den Burg, The Netherlands, and Centre for Ecological and Evolutionary Studies (CEES), University of Groningen, P.O. Box 14, 9750 AA Haren, The Netherlands