This research is supported by a Research Project Grant (RPG-097-2016) from the Leverhulme Trust. The Leverhulme Trust was established by the Will of William Hesketh Lever, the founder of Lever Brothers. Since 1925 the Trust has provided grants and scholarships for research and education. Today, it is one of the largest all-subject providers of research funding in the UK, distributing approximately £80m a year. For more information about the Trust, please visit www.leverhulme.ac.uk.