1. The Milner Centre for Evolution, Department of Biology & Biochemistry, University of Bath, Bath BA2 7AY, UK
2. Department of Life Sciences, The Natural History Museum, London SW7 5BD, UK
3. Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA
4. Department of Earth Sciences, University of Cambridge, Cambridge CB2 3EQ, UK