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''Atelopus varius'', the Costa Rican variable harlequin toad or clown frog, is a small
Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bi ...
true toad from the family
Bufonidae {{Cat main, Toad This category contains both species commonly called toads, and the true toads from the family Bufonidae. Animals by common name Frogs ...
(Crump 1986). Once ranging from Costa Rica to
Panama Panama ( , ; es, link=no, Panamá ), officially the Republic of Panama ( es, República de Panamá), is a transcontinental country spanning the southern part of North America and the northern part of South America. It is bordered by Co ...
, ''A. varius'' is now listed as critically endangered and has been reduced to a single remnant population near Quepos, Costa Rica (rediscovered in 2003) and has only relict populations in western Panama (IUCN). Recent variation in air temperature, precipitation, stream flow patterns, and the subsequent spread of a pathogenic
chytrid Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytri ...
fungus ''(
Batrachochytrium dendrobatidis ''Batrachochytrium dendrobatidis'' ( ), also known as ''Bd'' or the amphibian chytrid fungus, is a fungus that causes the disease chytridiomycosis in amphibians. Since its discovery in 1998 by Lee Berger, the disease devastated amphibian popula ...
)'' linked to global climate change have been the leading cause of decline for ''A. varius'' (Lips et al. 2003 and Pounds et al. 2006). '' A. zeteki'' has been considered a subspecies of ''A. varius'', but is now generally considered a separate species (Savage, 2002).


Geographic range

The historic range of A. varius stretched from the Pacific and Atlantic slopes of the Cordilleras de Tilaran mountain range in Costa Rica into western Panama. Suitable habitat includes both pre-montane and lower-montane zones as well as some lowland sites along rocky streams in hilly areas (ranging from 6 to 2000m in elevation) (Savage 1972).


Habitat and ecology

''A. varius'' is a diurnal frog often found on rocks or in crevices along streams in humid lowland and montane forests (Crump and Pounds 1985). It is primarily a
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
species, only entering the water during breeding season, relying on spray from streams for moisture (Pounds and Crump 1994). The Costa Rican variable harlequin frog is slow moving and often remains in the same area for long periods of time. The conspicuous or aposematic coloration of ''A. varius'' likely serves as a warning to potential predators of the toxicity of the frog's
integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, shell, germ or rind. Etymology The term is derived from ''integumentum'', which is Latin for "a covering". In a transferred, or ...
which contains
tetrodotoxin Tetrodotoxin (TTX) is a potent neurotoxin. Its name derives from Tetraodontiformes, an order that includes pufferfish, porcupinefish, ocean sunfish, and triggerfish; several of these species carry the toxin. Although tetrodotoxin was discove ...
, a potent
neurotoxin Neurotoxins are toxins that are destructive to nerve tissue (causing neurotoxicity). Neurotoxins are an extensive class of exogenous chemical neurological insultsSpencer 2000 that can adversely affect function in both developing and matur ...
(Crump and Pounds 1985). Its main food source is small
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s that are most abundant during the dry season (Crump 1988). The only known predator of ''A. varius'' is a parasitic
sarcophagid Sarcophagidae () are a family of flies commonly known as flesh flies. They differ from most flies in that they are ovoviviparous, opportunistically depositing hatched or hatching maggots instead of eggs on carrion, dung, decaying material, or op ...
fly ''(Notochaeta bufonivora)'' which deposits its larvae on the surface of the frog's thigh. The larvae then proceed to burrow inside the frog and eat it from within (Pounds and Crump 1987).


Conservation status

In recent decades, ''A. varius'' has become increasingly rare throughout its geographic range. The first incidence of its disappearance was recorded after a census conducted between 1990 and 1992 near Monteverde, Costa Rica revealed zero individuals where its population had previously peaked at 751 adults (Crump and Pounds 1985 and Pounds and Crump 1994). By 1996, ''A. varius'' was believed to be extinct throughout Costa Rica, however, in a 2019 assessment, two subpopulations were discovered in the country. In Panama, mass mortality has drastically reduced populations of ''A. varius'' in recent years and it was believed to be locally extinct (Lips 1999), however the species was recorded again in the 2019 assessment. Several theories related to changes in climatic patterns have been put forth to account for the rapid decline of ''A. varius''. A trend toward rising temperatures across the tropics in the late 1980s and early 1990s has been implicated in the declines of multiple
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia al ...
and amphibian species including several ''Atelopus'' spp. (Pounds et al. 1999). More recently, an observed global decline in amphibian species richness has been linked to an outbreak of the
pathogenic In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a ger ...
chytrid fungus ''
Batrachochytrium dendrobatidis ''Batrachochytrium dendrobatidis'' ( ), also known as ''Bd'' or the amphibian chytrid fungus, is a fungus that causes the disease chytridiomycosis in amphibians. Since its discovery in 1998 by Lee Berger, the disease devastated amphibian popula ...
'' (Pounds et al. 2006). This pathogen can be transmitted between individuals through shed skin cells and is known to infect
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ...
ized body surfaces where it can impair cutaneous respiration and
osmoregulation Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration ...
thus resulting in mortality (Pounds et al. 2006). Current attempts to preserve ''A. varius'' include a recently initiated captive breeding program as well as continued efforts to protect vital forest habitat (IUCN). In March 2021, it was announced that the species had been successfully bred in captivity outside of Panama for the first time.


References

*Crump, M.L. 1986. Homing and site fidelity in a Neotropical frog, ''Atelopus varius'' (Bufonidae). ''Copeia'' 1986(4): 1007-1009. *Crump, M.L. 1988. "Aggression in harlequin frogs: male-male competition and a possible conflict of interest between the sexes." ''Animal Behaviour'' 36(4): 1064-1077. *Crump, M.L., and J.A. Pounds 1985. "Lethal Parasitism of an Aposematic Anuran (''Atelopus varius'') by ''Notochaeta bufonivora'' (Diptera: Sarcophagidae)." ''Journal of Parasitology'' 71(5), 588-591. *Lips, K.R. 1999. Mass mortality and population declines of anurans at an upland site in western Panama. ''Conservation Biology'' 13: 117-125. *Lips, K.R., Green, D.E. and Papendick, R. 2003. Chytridiomycosis in wild frogs from southern Costa Rica. ''Journal of Herpetology'' 37: 215-218. *Pounds, J.A., Bustamante, M.R., Coloma, L.A., Consuegra, J.A., Fogden, M.P.L., Foster, P.N., La Marca, E., Masters, K.L., Merino-Viteri, A., Puschendorf, R., Ron, S.R., Sánchez-Azofeifa, G.A., Still, C.J. and Young, B.E. 2006. Widespread amphibian extinctions from epidemic disease driven by global warming. ''Nature'' 439: 161-167. *Pounds, J.A., and Crump, M.L. 1987. "Harlequin Frogs Along a Tropical Montane Stream: Aggregation and the Risk of Predation by Frog-Eating Flies." ''Biotropica'' 19(4): 306-309. *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: 72-85. *Pounds, J.A., Fogden, M.P.L., and J.H. Campbell 1999. "Biological response to climate change on a tropical mountain." ''Nature'' 398(6728): 611-615. *Savage, J.M. 1972. The harlequin frogs, genus ''Atelopus'', of Costa Rica, and western Panama. ''Herpetologica'' 28: 77-94. *Savage, J.M. 2002. ''The Amphibians and Reptiles of Costa Rica.'' University of Chicago Press, Chicago.


External links

*
IUCN Red List of All Threatened SpeciesThe Rainmaker ProjectAtelopus.com - Fighting to survive
{{Taxonbar, from=Q310885 varius Amphibians of Costa Rica Endemic fauna of Costa Rica Amphibians described in 1856 Taxa named by Eduard von Martens Taxa named by Hinrich Lichtenstein Critically endangered fauna of North America