1. Ecology and Evolutionary Biology, University of Adelaide, Adelaide, South Australia 5005, Australia
2. School of Biological Sciences, University of Queensland, St Lucia, Queensland 4072, Australia
3. Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta, Canada T0J 0Y0
4. Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity, Humboldt Universität zu Berlin, Invalidenstrasse 43, 10115 Berlin, Germany