1. School of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, Birmingham B15 2TT, UK
2. Centre for Palaeobiology Research, School of Geography, Geology and the Environment, University of Leicester, Leicester LE1 7RH, UK
3. Centre for Palaeobiology Research, School of Museum Studies, University of Leicester, Leicester LE1 7RF, UK
4. Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta, Canada T0J 0Y0