1. Department of Biology, National University of Ireland, Maynooth, Co. Kildare, Ireland; 2School of Biological Sciences, University of Bristol, Bristol BS8 1TQ, UK; 3Department of Life Sciences, The Natural History Museum, London SW7 5BD, UK; and 4Institute of Biological, Environmental & Rural Sciences, Aberystwyth University, Aberystwyth SY23 3FG, UK