1. 1The School of Environmental Sciences, University CoIlege Scarborough, Filey Road, Scarborough, North Yorkshire, YO11 3AZ, UK;, Email: grahams@ucscarb.ac.uk
2. 2The Institute of CeIl, Animal & Population Biology, The University of Edinburgh, West Mains Road, Edinburgh EH9 3JT, UK;, Email: j.deag@ed.ac.uk