Atelopus varius (Lichtenstein & Martens, 1856)

Painted frog, Clown frog, Veragoa stubfoot toad, Costa Rican variable harlequin toad, Talamancan harlequin frog



Taxonomy & Nomenclature



Conservation Status

Last record: 1996/7

Rediscovered in 2003

IUCN RedList status: Critically Endangered



Costa Rica & Panama (western)


Biology & Ecology










Original scientific description:

Lichtenstein, H., and von Martens, E. (1856). Nomenclator Reptilium et Amphibiorum Musei Zoologici Berolinensis. Namenverziechniss der in der zoologischen Sammlung der Königlichen Universität zu Berlin aufgestellten Arten von Reptilien und Amphibien nach ihren Ordnungen, Familien und Gattungen.  Berlin.

Other references:

Andreone, Franco et al. (2022). Threatened and extinct amphibians and reptiles in Italian natural history collections are useful conservation tools. Acta Herpetologica 17(1): 45-58. doi:10.36253/a_h-12349

Avery, Cecile. (2016). Comparison of Growth Rates in two Captive Bred Species of Atelopus (Anura; BUFONIDAE), at El Valle Amphibian Conservation Center. Independent Study Project (ISP) Collection. Paper 2390.

Barrio-Amorós, César L. and Abarca, Juan. (2016). Another surviving population of the Critically Endangered Atelopus varius (Anura: Bufonidae) in Costa Rica. Mesoamerican Herpetology 3(1): 128-134.

Coleman, Jeffrey L. (2020). Anthropogenic Drivers and Chytridiomycosis: Untangling the Disappearances of the Golden Toad and Costa Rican Variable Harlequin Toad and Addressing Amphibian Decline. Bulletin of the Chicago Herpetological Society 55(8): 157-162.

Crump, M. L. (1986). Homing and site fidelity in a Neotropical frog, Atelopus varius (Bufonidae). Copeia 1986: 1007-1009.

Crump, M. L., Hensley, F. R., Clark, K. L. (1992). Apparent decline of the golden toad: Underground or extinct? Copeia 1992(2): 413-420.

Crump, M. L. and Pounds, J. A. (1985). Lethal parasitism of an aposematic anuran (Atelopus varius) by Notochaeta bufonivora (Diptera: Sarcophagidae). J. Parasitol. 71: 588-591.

Gómez-Hoyos, Diego A. et al. (2018). Phytotelmata selection by anurans and implications for their conservation at Las Tablas Protected Zone, Costa Rica. Alytes 35(1-4): 1-11.

Gómez–Hoyos, Diego A. et al. (2020). Potential effect of habitat disturbance on reproduction of the critically endangered harlequin frog Atelopus varius in Las Tablas, Costa Rica. Animal Biodiversity and Conservation 43(1): 1-7.

José F. González-Maya, Jerrold L. Belant, Sarah A. Wyatt, Jan Schipper, Josué Cardenal, Daniel Corrales, Iván Cruz-Lizano, Annelie Hoepker, Armando H. Escobedo-Galván, Fernando Castañeda and Addison Fischer. (2013). Renewing hope: the rediscovery of Atelopus varius in Costa Rica. Amphibia-Reptilia 34(4): 573-578. [automatic download]

González-Maya, José F., et al. (2019a). Depredacion por el Chinche de Agua (Abedus sp.; Belostomatidae) sobre el Criticamente Amenazado Atelopus varius (Bufonidae) en la Zona Protectora Las Tablas, Costa Rica. Revista Acta Biologica Colombiana 24(2): 403-406. [automatic download]

González-Maya, José F., et al. (2019b). WATER-BUG (Abedus SP.; BELOSTOMATIDAE) PREDATION ON THE CRITICALLY ENDANGERED Atelopus varius (BUFONIDAE) AT LAS TABLAS PROTECTED ZONE, COSTA RICA. Acta Biológica Colombiana 24(2).

Hertz, Andreas et al. (2012). Field notes on findings of threatened amphibian species in the central mountain range of western Panama. Amphibian and Reptile Conservation 6(2): 9-30.

Jaynes, Kyle E. et al. (2022). Harlequin frog rediscoveries provide insights into species persistence in the face of drastic amphibian declines. Biological Conservation 276: 109784.

Jiménez-Monge A, Montoya-Greenheck F, Bolaños F, Alvarado G. 2019. From incidental findings to systematic discovery: locating and monitoring a new population of the endangered Harlequin Toad. Amphibian & Reptile Conservation 13(2): [General Section]: 115–125 (e192).

La Marca, E., Lips, K. R., Lötters, S., Puschendorf, R., Ibáñez, R., Rueda-Almonacid, J. V., Schulte, R., Marty, C., Castro, F., Manzanilla-Puppo, J., García-Pérez, J. E., Bolaños, F., Chaves, G., Pounds, J. A., Toral, E. and Young, B. E. (2005). Catastrophic population declines and extinctions in neotropical Harlequin frogs (Bufonidae: Atelopus). Biotropica 37(2): 190-201.

Lips, K. R. (1998). Decline of a tropical montane amphibian fauna. Conservation Biology 12(1):106-117.

Lips, K. R. (1999). Mass mortality and population declines of anurans at an upland site in western Panama. Conservation Biology 13(1): 117-125.

Lips, K. R., Reeve, J. D. and Witters, L. R. (2003). Ecological traits predicting amphibian population declines in Central America. Conservation Biology 17(4): 1078-1088.

McCaffery, Rebecca et al. (2015). The demography of Atelopus decline: Harlequin frog survival and abundance in central Panama prior to and during a disease outbreak. Global Ecology and Conservation 4: 232-242.

Perez R, Richards-Zawacki CL, Krohn AR, Robak M, Griffith EJ, Ross H, Gratwicke B, Ibáñez R, Voyles J. (2014). Field surveys in Western Panama indicate populations of Atelopus varius frogs are persisting in regions where Batrachochytrium dendrobatidis is now enzootic. Amphibian & Reptile Conservation 8(2) [General Section]: 30-35 (e85).

Pounds, J. A. and Crump, M. L. (1994). Amphibian declines and climate disturbance: The case of the golden toad and the harlequin frog. Conservation Biology 8(1): 72-85.

Alan Pounds, Robert Puschendorf, Federico Bolaños, Gerardo Chaves, Martha Crump, Frank Solís, Roberto Ibáñez, Jay Savage, César Jaramillo, Querube Fuenmayor, Karen Lips. (2010). Atelopus varius. In: IUCN 2013. IUCN Red List of Threatened Species. Version 2013.2. ( Downloaded on 24 November 2013.

Ramírez, Juan P. et al. (2020). Recent and Rapid Radiation of the Highly Endangered Harlequin Frogs (Atelopus) into Central America Inferred from Mitochondrial DNA Sequences. Diversity 12(9): 360.

Richards-Zawacki, C. L. (2009). Effects of slope and riparian habitat connectivity on gene flow in an endangered Panamanian frog, Atelopus varius. Diversity and Distributions15: 796-806.

Ryan, M., Berlin, E. and Gagliardo, R. (2005). Further exploration in search of Atelopus varius in Costa Rica. FrogLog 69: 1-2.

Scheffers, Brett R., Yong, Ding Li, Harris, J. Berton C., Giam, Xingli and Sodhi, Navjot S. (2011). The world’s rediscovered species: back from the brink? PLoS ONE 6(7): e22531. [Supporting Information (Table S1)];hitNum=1#page-2


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