[1] Univ Tours, Inst Rech Biol Insecte, UMR 7261, CNRS, F-37200 Tours, France
[2] INRA, Ctr Rech Provence Alpes Cote Azur, UR Abeilles & Environm 406, Site Agroparc,Domaine St Paul 228, F-84914 Avignon, France
[3] Western Sydney Univ, Hawkesbury Inst Environm, Locked Bag 1797, Penrith, NSW 2751, Australia
[4] Chinese Acad Agr Sci, Inst Apicultural Res, Beijing 100093, Peoples R China
[5] Minist Agr, Key Lab Pollinating Insect Biol, Beijing 100093, Peoples R China
[6] Univ Montpellier, CNRS, IRD, EPHE,Inst Sci Evolut,UMR5554, Montpellier, France
[7] Univ Cambridge, Dept Vet Med, Madlngley Rd, Cambridge CB3 0ES, England