[1] Inst Nacl de Pesquisas da Amazonia, Posgrad Ecol, Manaus, Brazil
[2] Univ Estadual Mato Grosso do Sul, Lab Bioecol Scarabaeoidea Scaralab, Aquidauana, Brazil
[3] Univ Fed Para, Sinteses Biodiversidade Amazon INCT SinBiAm, Belem, Brazil
[4] Univ Fed Amazonas, Dept Biol, Manaus, Brazil
[5] Univ Pretoria, Dept Zool & Entomol, Intertebrate Systemat & Conservat Grp, Hatfield, South Africa
[6] Univ Fed Mato Grosso UFMT, Lab Scarabaeoidol, Inst Biociencias, Cuiaba, Brazil