1 Department of Biological Sciences and Institute for Bioinformatics and Evolutionary Studies, University of Idaho, Moscow, ID 83844, 2 Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, 3 Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI 48109, USA, 4 Department of Biological Sciences, Macquarie University, Sydney, NSW 2109, Australia and 5 Department of Ecology and Evolutionary Biology, University of California, Los Angeles, Los Angeles, CA 90095, USA (At 2014-04-10)