[1] Univ Nacl, Escuela Ciencias Biol, Heredia 863000, Costa Rica
[2] John Carroll Univ, Dept Biol, University Hts, OH 44118 USA
[3] Univ Costa Rica, Escuela Biol, San Jose 115012060, Costa Rica
[4] Univ Costa Rica, Inst Clodomiro Picado, San Jose 115012060, Costa Rica