Institute of Evolutionary Biology, University of Edinburgh, Kings Buildings, West Mains Road, EH9 3JT, UK. jerome.kelleher@ed.ac.uk|Institute of Evolutionary Biology, University of Edinburgh, Kings Buildings, West Mains Road, EH9 3JT, UK. jerome.kelleher@ed.ac.uk|Institute of Evolutionary Biology, University of Edinburgh, Kings Buildings, West Mains Road, EH9 3JT, UK. jerome.kelleher@ed.ac.uk