[1] Univ Austral Chile, Fac Ciencias Forestales & Recursos Nat, Inst Conservac Biodiversidad & Terr, Valdivia, Chile
[2] McGill Univ, Dept Biol, Montreal, PQ H3A 1B1, Canada
[3] McGill Univ, Redpath Museum, Montreal, PQ H3A 2K6, Canada
[4] Smithsonian Trop Res Inst, Panama City, Panama
[5] Univ Austral Chile, Fac Ciencias, Inst Ciencias Ambientales & Evolut, Valdivia, Chile