[1] Univ Hertfordshire, Landscape & Ecol Res Grp, Dept Environm Sci, Hatfield AL10 9AB, Herts, England
[2] Univ New England, Biodivers & Conservat Unit, Dept Land & Water Conservat, Armidale, NSW 2350, Australia
[3] Univ Plymouth, Environm Sci Lab, Fac Sci, Plymouth PL4 8AA, Devon, England
[4] Chester Coll, Dept Geog, Chester CH1 4BJ, Cheshire, England