1. Department of Zoology, Downing Street, Cambridge, CB2 3EJ; 2Division of Ecology and Evolution, Imperial College London, South Kensington Campus, London, SW7 2AZ; 3Department of Life Sciences, Natural History Museum, Cromwell Road, London, SW7 5BD; and 4Department of Physics, Cavendish Laboratory, JJ Thomson Avenue, Cambridge, CB3 0HE, UK