1. Department of Zoology, University of Western Australia, Nedlands, WA 6907,Australia
2. Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK
3. Author for correspondence (e-mail: henry.bennet-clark@zoo.ox.ac.uk)
4. Department of Electronic Materials, Engineering Research School of Physical Sciences and Engineering, Australian National University, Canberra,ACT 0200, Australia