1. Program of Population Biology, Ecology, and Evolution, Emory University, Atlanta, GA, 30322, USA
2. Biodiversity and Tropical Ecology Research Center (CIBET), School of Biology, University of Costa Rica, 11501-2060 San José, Costa Rica
3. CIET-Virology, Microbiology, University of Costa Rica, 11501-2060 San José, Costa Rica
4. Department of Wildlife Diseases, Leibniz Institute for Zoo and Wildlife Research, Alfred-Kowalke-Str. 17, 10315 Berlin, Germany
5. Program of Population Biology, Ecology, and Evolution, Emory University, Atlanta, GA, 30322, USA; Biology Department, Emory University, Atlanta, GA, 30322, USA
6. Program of Population Biology, Ecology, and Evolution, Emory University, Atlanta, GA, 30322, USA; Department of Environmental Sciences, Emory University, Atlanta, GA, 30322, USA