[1] RSPB, Ctr Conservat Sci, Sandy SG19 2DL, Beds, England
[2] Univ Aberdeen, Sch Biol Sci, Inst Biol & Environm Sci, 23 St Machar Dr, Aberdeen AB24 3UU, Scotland
[3] RSPB Scotland, Ctr Conservat Sci, 2 Lochside View,Edinburgh Pk, Edinburgh EH12 9DH, Midlothian, Scotland
[4] Conservat Sci Grp, David Attenborough Bldg,Pembroke St, Cambridge CB2 3QZ, England