1. Ecosystems Department, British Antarctic Survey, Madingely Road, Cambridge, CB3 OET, UK
2. Fisheries Ecology Division, Southwest Fisheries Science Center, National Marine Fisheries Service, National Oceanic and Atmospheric Administration, 110 McAllister Way, Santa Cruz, CA 95060, USA
3. Department of Applied Mathematics, University of California Santa Cruz, 110 McAllister Way, Santa Cruz, CA 95060, USA
4. School of Ocean Sciences, Bangor University, Isle of Anglesey LL59 5AB, UK