[1] Univ Costa Rica, Inst Clodomiro Picado, San Jose, Costa Rica
[2] Univ Costa Rica, Ctr Invest Biodiversidad & Ecol Trop, Museo Zool, San Jose, Costa Rica
[3] Univ Valencia, Inst Cavanilles Biodiversidad & Biol Evolut, Valencia, Spain
[4] Org Estudios Trop, San Jose, Costa Rica