[1] Royal Holloway Univ London, Ctr Ecol Evolut & Behav, Sch Life Sci & Environm, Dept Biol Sci, Egham, England
[2] Imperial Coll London, Dept Life Sci, London, England
[3] UCL, Ctr Ecol & Evolut, London, England
[4] Zool Staatssammlung, Hymenoptera Sect, Munich, Germany
[5] Natl Museums Scotland, Dept Nat Sci, Edinburgh, Scotland
[6] Nat Hist Museum, London SW7 5BD, England
[7] Stirling Univ, Beta Ctr, Bumblebee Conservat Trust, Innovat Pk, Stirling, Scotland