Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK. keith.jolley@ceid.ox.ac.uk|Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK. keith.jolley@ceid.ox.ac.uk|Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK. keith.jolley@ceid.ox.ac.uk|Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK. keith.jolley@ceid.ox.ac.uk