This work was funded by the UK Natural Environment Research Council (through grants to Nienow, Mair, and Wadham, and a studentship to Bartholomew), the Edinburgh University Moss Centenary Scholarship (Cowton and Bartholomew), and a Carnegie Research Grant (Nienow). We thank Ian Willis, Tim Bartholomaus and an anonymous referee for valuable comments which significantly improved the manuscript.Peer reviewedPublisher PD