[1] Open Univ, Environm Earth & Ecosyst, Walton Hall, Milton Keynes MK7 6AA, Bucks, England
[2] Florida Inst Technol, Biol Sci, 150 West Univ Blvd, Melbourne, FL 32901 USA
[3] Univ Amsterdam, IBED, Palaeoecol & Landscape Ecol, Sci Pk 904, NL-1098 XH Amsterdam, Netherlands
[4] Univ Reading, Dept Geog & Environm Sci SAGES, Ctr Past Climate Change, Reading RG6 6AB, Berks, England
[5] Northwestern Univ, Dept Earth & Planetary Sci, 2145 Sheridan Rd, Evanston, IL 60201 USA
[6] Nat Hist Museum, Dept Life Sci, Cromwell Rd, London SW7 5BD, England