1. British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley Road, Cambridge CB3 0ET, UK
2. Department of Biosciences, Durham University, South Road, Durham DH1 3LE, UK
3. Woodwell Climate Research Center, Woods Hole Road, Falmouth, MA 02540, USA
4. Institute of Arctic Biology, University of Alaska Fairbanks, Koyukuk Drive, Fairbanks, AK 99775, USA