[1] Univ Tennessee, Dept Ecol & Evolutionary Biol, Knoxville, TN 37996 USA
[2] Univ Sheffield, Biodivers & Macroecol Grp, Dept Anim & Plant Sci, Sheffield S10 2TN, S Yorkshire, England
[3] Mapping GIS & Spatial Solut, Sheffield, S Yorkshire, England
[4] Yorkshire Wildlife Trust, York YO24 1GN, N Yorkshire, England