[1] Univ Sussex, Sch Life Sci, Lab Apiculture & Social Insects, Brighton, E Sussex, England
[2] Univ Sao Paulo, Luiz de Queiroz Coll Agr, Dept Entomol & Acarol, Piracicaba, Brazil
[3] Univ Sao Paulo, Biosci Inst, Dept Zool, Sao Paulo, Brazil
[4] Calif Dept Food & Agr, Plant Pest Diagnost Ctr, Sacramento, CA 95814 USA
[5] Univ Florida, Dept Entomol & Nematol, Honey Bee Res & Extens Lab, Gainesville, FL 32611 USA