1. Biological and Environmental Sciences, Faculty of Natural Sciences, University of Stirling , FK9 4NF , UK
2. Laboratory of Genetics, Plant Sciences Group, Wageningen University and Research , 6708 PB , The Netherlands
3. Charles Darwin Research Station, Charles Darwin Foundation , Santa Cruz, Galapagos Islands , Ecuador
4. Galapagos National Park Directorate , Santa Cruz, Galapagos Islands , Ecuador