[1] Univ Sao Paulo MZSP, Museu Zool, Lab Coleoptera LaC, Sao Paulo, SP, Brazil
[2] Univ Sao Paulo, Fac Filosofia Ciencias & Letras Ribeirao Preto FF, Dept Biol, Programa Posgrad Entomol, Ribeirao Preto, SP, Brazil
[3] Nat Hist Museum, Dept Life Sci, London SW7 5BD, England
[4] Univ Estadual Paulista Julio de Mesquita Filho Un, Fac Ciencias & Letras Assis FCL, Dept Ciencias Biol, Lab Biol Aquat LABIA, Assis, SP, Brazil