Main Banner
Browse Species

Atelopus varius Lichtenstein and von Martens 1856

Critically Endangered (IUCN 3.1)
Atelopus varius
Male Harlequin Frog (Atelopus varius)

Common name

Harlequin Frog, Clown Frog, Painted Frog, Veragoa Stubfoot Toad


Atelopus varius has experienced precipitous population declines throughout its range. Populations west of the Canal in Panama have shown drastic declines, and it is nearly extinct in Costa Rica. Chytrid fungus is thought to be the primary cause of its disappearance.



Species description based on Savage (2002). Medium frog (males 27-39 mm, females 33-48 mm).

Atelopus varius Adult 1


Dorsal pattern and coloration extremely variable, but generally consisting of black and one of many colors: bright green, yellow, yellow-orange, or red. Savage (1972) has some excellent photographs showing the range of variation in patterning in this species. The dorsal surface is mostly smooth and lacks any glandular concentrations apparent in some other species of Atelopus.

Atelopus varius Dorsal 1 Atelopus varius Dorsal 2


Ventral surface usually yellow, sometimes mixed with green or red.

Atelopus varius Ventral 1


Iris green. Pupil horizontal.

Life history

Breeding season

In Monteverde, Costa Rica, oviposition takes place during October-December (Savage 2002). In Sante Fe, Panama, an amplectant pair was found in June, but only non-calling males were observed in July (CoCroft et al 1990).

Atelopus varius Breeding season 1 Atelopus varius Breeding season 2


Eggs are laid in two strings and are white (Starret 1967). Eggs are probably affixed to rocks to prevent them from flowing downstream (Starrett 1967). Eggs hatch 6 days after oviposition (Starrett 1967).


The dark brown body is very small, rather flat and looks square from above (Savage 2002). The mouth is very large (Savage 2002). The ventral surface has a large disk that likely serves as a sucker to help tadpoles adhere to rocks in fast-moving water (Savage 2002, Duellman and Lynch 1969). An excellent illustration of the tadpole can be found in Starrett (1967). Tooth rows are 2/3 (Starrett 1967).

Ecology behavior and evolution


Atelopus varius is an ant specialist, but feeds in small amounts on other arthropods (Toft 1981). They actively search for their prey (Toft 1981).


This species is often encountered on rocks in or along the margins of streams (Crump 1986). At night, they sleep on rocks or vegetation (Crump 1986). Population declines have been documented throughout Costa Rica and parts of Panama (Pounds and Crump 1994). Adult Atelopus varius sometimes become parasitized by flies, which leads to the death of the frog (Crump and Pounds 1985). Frogs are more vulnerable to parasitism during the dry season, when they are forced to aggregate in suitable waterfall spray habitat near streams (Pounds and Crump 1987, Crump and Pounds 1989).


A buzz repeated numerous times (Savage 2002). Atelopus varius also produces chirps that may serve as release calls (Savage 2002). An audiospectrogram of the call can be found in Cocroft et al. (1990).

Behavior and communication

Atelopus varius moves very slowly; presumably the toxins in their skin protect them from potential predators (Crump 1986). During the breeding season, males call to defend territories, but aggressive interactions may escalate to wrestling to establish dominance (Savage 2002). Females are more territorial during the non-breeding season (Savage 2002). This species exhibits homing behavior, returning to the place of capture within a week or less of displacement (Crump 1986).


2N = 22 (Duellman 1967, Schmid 1980)


The skin of Atelopus varius contains tetrodotoxin (Kim et al. 1975).

Taxonomy and systematics


  • Kingdom:Animalia


Lichtenstein and von Martens 1856


Atelopus bibronii, Atelopus loomisi, Atelopus varius ambulatorius, Atelopus varius bibroni, Atelopus varius loomisi, Atelopus varius maculatus, Atelopus varius varius, Atelopus varius var. maculatus, Hylaemorphus bibronii, Hylaemorphus dumerilii, Hylaemorphus pluto, Hylaemorphus plutonius, Phrynidium varium, Phrynidium varium var. adspersum, Phrynidium varium var. maculatum, Phryniscus bibronii, Phryniscus dumerilii, Phryniscus varius

Type locality

"Veragoa". Restricted to "the Pacific portion of the Provincia Veraguas, western Panamá" by Lotters et al (1998)

Habitat and distribution


Mostly found in premontane and montane forest to 2000 m but also on hills in some lowland sites.


Panama. This species has almost gone extinct in Costa Rica.


Atelopus varius distribution
Distrubution map (IUCN)


Berger, L, R Speare, P Daszak, DE Green, AA Cunningham, CL Goggin, R Slocombe, MA Ragan, AD Hyatt, KR McDonald, HB Hines, KR Lips, G Marantelli, and H Parkes. 1998. Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America. PNAS 95(15): 9031-9036.

Cocroft, RB, RW McDiarmid, AP Jaslow and PM Ruiz-Carranza. 1990. Vocalizations of eight species of Atelopus (Anura: Bufonidae) with comments on communication in the genus. Copeia 1990(3): 631-643.

Crump, ML and JA Pounds. 1985. Lethal parasitism of an aposematic anuran (Atelopus varius) by Notochaeta bufonivora (Diptera: Sarcophagidae). Journal of Parasitology 71(5): 588-591.

Crump, ML and JA Pounds. 1989. Temporal variation in the dispersion of a tropical anuran. Copeia 1989(1): 209-11.

Crump, ML. 1986. Homing and site fidelity in a Neotropical frog, Atelopus varius (Bufonidae). Copeia 1986(2): 1007-1009.

Crump, ML. 1988. Aggression in Harlequin frogs: male-male competition and a possible conflict of interest between the sexes. Animal Behaviour 36(4): 1064-1077.

Daly, JW, F Gusovsky, CW Myers, M Yotsu-Yamashita, and T Yasumoto. 1994. First occurrence of tetrodotoxin in a dendrobatid frog (Colostethus inguinalis), with further reports for the bufonid genus Atelopus. Toxicon 32(3): 279-85.

Daly, JW, WL Padgett, RL Saunders, and JF Cover, Jr. 1997. Absence of tetrodotoxins in a captive-raised riparian frog, Atelopus varius. Toxicon 35(5): 705-709.

Duellman, WE. 1967. Additional Studies of Chromosomes of Anuran Amphibians. Systematic Zoology 16(1): 38-43.

Dunn, ER. 1940. New and Noteworthy Herpetological Material from Panama. Proceedings of the Academy of Natural Sciences of Philadelphia 92: 105-122.

Emerson, SB. 1978. Allometry and Jumping in Frogs: Helping the Twain to Meet. Evolution 32(3): 551-564.

Emerson, SB. 1982. Frog Postcranial Morphology: Identification of a Functional Complex. Copeia 1982(3): 603-613.

Erspamer, V, G Erspamer Falconieri, and JM Cei. 1986. Active peptides in the skins of two hundred and thirty American amphibian species. Comparative Biochemistry and Physiology. C. Comparative pharmacology 85(1): 125-137.

Hailman, JP and RG Jaeger. 1974. Phototactic responses to spectrally dominant stimuli and use of colour vision by adult anuran amphibians: A comparative survey. Animal Behaviour 22(4): 757-795.

Howland, HC, M Howland, A Giunta and TW Cronin. 1997. Corneal curvatures and refractions of central American frogs. Vision Research37(2): 169-174.

Ibáñez, R, F Solís, C Jaramillo, and S Rand. 2000. An overwiew of the herpetology of Panama. In: Johnson, JD, RG Webb, and OA Flores-Villela. Eds. Mesoamerican Herpetology: Systematics, Zoogeography and Conservation. The University of Texas at El Paso, El Paso, Texas.

Kim, YH, GB Brown, HS Mosher, and FA Fuhrman. 1975. Tetrodotoxin: occurrence in atelopid frogs of Costa Rica. Science 189: 151-152.

Kim, YH, Y-B Kim, and M Yotsu-Yamashita. 2003. Potent neurotoxins: Tetrodotoxin, Chiriquitoxin, and Zetekitoxin from Atelopus frogs in Central America. Journal of Toxicology: Toxin Reviews 22(4): 521-532.

La Marca, E, KR Lips, S Lötters, R Puschendorf, R Ibáñez, JV Rueda-Almonacid, R Schulte, C Marty, F Castro, J Manzanilla-Puppo, JE Garcia-Perez, E Toral, F Bolaños, G Chaves, JA Pounds, and B Young. 2005. Catastrophic population declines and extinctions in Neotropical harlequin frogs (Bufonidae: Atelopus). Biotropica 37(2)190-201.

Lips, K and JM Savage. 1996. Key to the Known Tadpoles (Amphibia: Anura) of Costa Rica. Studies on Neotropical Fauna and Environment 31(1): 17-26

Lips, KR, DE Green, and R Papendick. 2003. Chytridiomycosis in wild frogs from southern Costa Rica. Journal of Herpetology 37(1): 215-218.

Lips, KR, JD Reeve, and LR Witters. 2003. Ecological traits predicting amphibian population declines in Central America. Conservation Biology 17(4): 1078-1088.

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

Lötters, S, W Böhme 2, and R Günther. 1998. Notes on the type material of the Neotropical Harlequin Frogs Atelopus varius (Lichtenstein & Martens, 1856) and Atelopus cruciger (Lichtenstein & Martens, 1856) deposited in the Museum für Naturkunde of Berlin (Anura, Bufonidae). Zoosystematics and Evolution 74(2): 173-184.

Lötters, S. 1996. The Neotropical Toad Genus Atelopus. Checklist - Biology - Distribution. Vences, M and F Glaw. Eds. Verlags GbR, Köln, Germany.

Navas, CA. 1996. Metabolic physiology, locomotor performance, and thermal niche breadth in neotropical anurans. Physiological Zoology 69:1481-1501.

Pounds, JA and ML Crump. 1987. Harlequin Frogs along a tropical montane stream: aggregation and the risk of predation by frog-eating flies. Biotropica 19(4): 306-309.

Pounds, JA and ML Crump. 1989. Temporal variation in the dispersion of a tropical anuran. Copeia 1989(1): 209-211.

Pounds, JA and ML Crump. 1994. Amphibian declines and climate disturbance: the case of the Golden Toad and the Harlequin Frog. Conservation Biology 8(1): 72-85.

Pounds, JA, MPL Fogden, and JH Campbell. 1999. Biological response to climate change on a tropical mountain. Nature 398(6728): 611-615.

Pounds, JA, MR Bustamante, LA Coloma, JA Consuegra, MPL Fogden, PN Foster, E La Marca, KL Masters, A Merino-Viteri, R Puschendorf, SR Ron, GA Sánchez-Azofeifa, CJ Still, and BE Young. 2006. Widespread amphibian extinctions from epidemic disease driven by global warming. Nature 439: 161-167.

Puschendorf, R, F Bolanos, and G Chaves. 2006. The amphibian chytrid fungus along an altitudinal transect before the first reported declines in Costa Rica. Biological Conservation 132(1): 136-142.

Puschendorf, R. 2003.Atelopus varius (Harlequin Frog) fungal infection. Atelopus varius (Rana arlequín) infección fungosa. P.imprenta: v. 34, no. 4, p. 355. Año 2003.

Richards-Zawacki, CL. 2009. Effects of slope and riparian habitat connectivity on gene flow in an endangered Panamanian frog, Atelopus varius. Diversity and Distributions 15(5): 796-806.

Savage, JM. 1972. The harlequin frogs, genus Atelopus, of Costa Rica, and western Panama. Herpetologica 28(2): 77-94.

Savage, JM. 2002. The Amphibians and Reptiles of Costa Rica: A Herpetofauna between two Continents, between two Seas. University of Chicago Press, Chicago.

Starrett, P. 1967. Observations on the life history of frogs of the family Atelopodidae. Herpetologica 23(3): 195-204.

Taylor, EH. 1952. A review of the frogs and toads of Costa Rica. Revisión de las ranas y sapos de Costa Rica. The University of Kansas Science Bulletin: 35(1): 577-941.

Toft, CA. 1981. Feeding ecology of Panamanian litter anurans: patterns in diet and foraging mode. Herpetologica 15(2): 139-44.

Woodhams, DC, J Voyles, KR Lips, C Carey and LA Rollins-Smith. 2006. Predicted disease susceptibility in a Panamanian amphibian assemblage based on skin peptide defenses. Journal of Wildlife Diseases 42(2): 207-218.

Yeh, J. 2002. The effect of miniaturized body size on skeletal morphology in frogs. Evolution 56(3): 628-641.

Additional resources