[1] Univ York, Hull York Med Sch, York YO10 5DD, England
[2] UCL, Ctr Integrat Anat, Dept Cell & Dev Biol, London WC1E 6BT, England
[3] Univ Reading, Dept Geog & Environm Sci, Reading RG6 6DW, England
[4] Natl Museums Scotland, Dept Nat Sci, Edinburgh EH1 1JF, Scotland
[5] Univ Edinburgh, Inst Geog, Sch Geosci, Edinburgh EH8 9XP, Scotland
[6] Nottingham Trent Univ, Sch Anim Rural & Environm Sci, Nottingham NG25 0QF, England
[7] Univ Exeter, Dept Archaeol & Hist, Exeter EX4 4QJ, England