1. Department of Ecology & Evolutionary Biology, Cornell University, 215 Tower Road, Ithaca NY, 14853, USA;
2. Department of Geology & Geophysics, Yale University, 210 Whitney Avenue New Haven, CT, 06511, USA;
3. Department of Biology and Biochemistry, Milner Centre for Evolution, University of Bath, Building 4 South, Claverton Down, Bath, BA2 7AY, UK