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adult counterparts, the juveniles go to higher elevations and recede into trees to prevent predation; however, due to their small size, the juveniles are not able to cover as much elevation and climb as highly into trees and shrubbery. At the onset of heavy and consistent rains, the juveniles flee from the open streamsides, which are normally where the grown adult males, which are very territorial, are known to roam. Territorial behaviors by adult males could be initiated by these rains. Visually, the juvenile has snout-to-vent lengths ranging from 8.4 to 17.1 mm. Their dorsal coloration is a deep and vivid green which matches the color of the moss that grows on the rocks in and around the streams of their habitat. There are also dark brown to black dorsal markings. Some of the juveniles are also known to have small dark markings on their digits. Their abdomen is either white or goldenrod yellow, occasionally with dark markings that do not match the ground color.
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pathogens. Researchers characterized the bacterial community from wild and captive
Panamanian golden frogs originating from the same population with sequencing to assess how long-term captivity has affected this community. Species richness, phylogenetic diversity, and community structure of the skin microbiota were found to be significantly different between wild and captive Panamanian golden frogs. However, after approximately eight years of living in captivity, the offspring of the original captive Panamanian golden frogs still shared 70% of their microbial community with wild frogs. These results demonstrate that host-associated microbial communities can be significantly altered by captive management, but most of the community composition can be preserved.
531:, are able to metabolize the frogs' poison. Ways to ward off predators and prevent predation are different in their diurnal habits versus their nocturnal ones, especially because the poison alone will not ward off every predator. Adult males, which are active on the ground during the day, recede into the trees and perch there at night. This is most likely a defense mechanism. If the predator is approaching at night, the frog cannot rely on a visual strategy for fleeing. They perch on trees because it gives them the advantage of hearing approaching predators or feeling their weight on the tree branch. The noise and tactile advantages of climbing a tree are better than the advantages of burrowing in the ground.
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654:. When the toad dies, it is thought to turn to gold and to bring good luck to those fortunate enough to see it. In 2010, the Panamanian government passed legislation recognizing August 14 as "National Golden Frog Day". The main celebration event is marked annually by a parade in the streets of El Valle de Anton, and a display of Panamanian golden frogs at the El Valle Amphibian Conservation Center in the El Nispero Zoo, El Valle. The highly toxic skin of the frog has also been used for centuries by the native people of the Panamanian forests for arrow poison.
468:. They are typically around 5.8 mm in length and 4.3 mm in width. Their snouts are rounded, as well as their tails. The longest caudal fins on their tails are about three-fifths the length of the tail. Their mouths are large and ventral surrounded by labia which form an unbroken oral disc about 3.6 mm wide. The posterior lip has no papillae, but other lips are lined with single rows of small, blunt papillae. They are colored from dark brown to black dorsally, with golden flecks on their bodies. They develop this black and gold coloration as
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inhibit its ability to communicate with other members of its species through throat sounds. Despite lacking eardrums, the "earless" frog responds to vocalizations produced by members of its own species. The male frog responds to a pulsed vocalization, characterized by lower frequencies followed by higher frequencies, and so on, by exhibiting antagonistic behaviors such as turning to face the source of the sound and producing a pulsed vocalization in response. The pulsed call is used to demonstrate male position during combative situations. Like
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governmental agencies and nongovernmental organizations, heightened awareness of current global amphibian declines, greater respect for wildlife among
Panamanians and global citizens, and greater land preservation for threatened and endangered species throughout the world. This organization will use education and field studies, producing offspring through the already captive frogs, and offering financial support to help preserve these frogs.
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510:, it is very territorial, living in the same site most of its life. As a result of this site fidelity, it will not hesitate to vocalize when another male frog encroaches on its territory. If this is not enough to get the intruder away, the frog is not hesitant to defend its territory through aggressive behavior. When encountering another male, male frogs will wave their forefeet as a sign of defense.
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456:, juvenile, subadult, and adult. During the larval stage, individuals emerge from their eggs after 2 to 10 days of development. They are entirely aquatic creatures at this stage and are found in waters with a temperature range from 20.4 to 21.3 °C and with depths of 5 to 35 cm. After emerging, they mostly spend time resting in shallow pools below cascades. This behavior is similar to
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fungus, their body temperatures rise to fight off the fungus. However, even if the infection leaves the frogs and body temperatures return to previous normal levels, the infection can re-emerge. Another study found that dry conditions added an average 25 days to the lifespan of infected individuals, while higher temperatures only added 4 days.
400:, which caused the El Copé population to rapidly collapse in 2004. Vital habitat is lost each year to small farms, commercialized agriculture, woodlot operations, livestock range, industrial expansion, and real estate development. Individuals are kept in captive-breeding programmes in more than 50 institutions across North America and Panama.
368:. Their toxin is water-soluble and affects the nerve cells of anyone who comes in contact with it. Panamanian golden frogs use this toxin to protect themselves from most predators. Due to the risk of testing the poison on humans, it has been done with mice. Large doses can be fatal in 20 or 30 minutes. Death is preceded by
554:. Although captive populations seem to thrive well, reintroducing them to an area will not stop the threat of chytridiomycosis. No current remedies prevent or control the disease in the wild, but efforts are being made. One attempt was made to protect a wide variety of frogs from the disease by using the bacterium
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encounters a predator, it often waves and lifts its foot at the predator to call attention to its stunning and beautiful coloring. This coloring is a warning of its toxicity, which is enough to make a predator no longer consider the frog as a meal. If the predator continues to approach, undeterred by
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The
Panamanian golden frog appears to socialize with other amphibians using sounds from the throat and hand-waving, like the semaphore motion used in courtship. The movements may be friendly or aggressive warnings. It is an "earless" species of frog, meaning it lacks tympanum. This, however, does not
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The subadults of this species have full ranges, but they are sometimes found near adult males which is noteworthy because males of this species are fairly solitary and combative in the presence of other males. The subadult is about 28.3 mm long and weights about 1.1 g. They are more greenish in
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in the
Panamanian golden frog because of the noise of the fast-moving streams which formed its natural habitat. The male tends to stay near the streams where breeding occurs, while in the nonbreeding season, the female retreats into the forests. The male uses a soft call to entice prospective mates,
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The
Panamanian golden frog, apart from recognizing sounds, is also able to locate the origin of a sound. This means it is capable of directional hearing. In all other species of frogs, the role of the eardrum is to pinpoint the direction of sound. Due to the very small size of the Panamanian golden
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Becker, Matthew H.; Harris, Reid N.; Minbiole, Kevin P. C.; Schwantes, Christian R.; Rollins-Smith, Louise A.; Reinert, Laura K.; Brucker, Robert M.; Domangue, Rickie J.; Gratwicke, Brian (2012). "Towards a better understanding of the use of probiotics for preventing chytridiomycosis in
Panamanian
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fungus into
Panamanian golden frog habitat, conservation organizations collected Panamanian golden frogs and placed them in captive survival assurance colonies. The skin of amphibians is host to a diverse resident bacterial community, which acts as a defense mechanism in some amphibians to inhibit
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started a conservation effort and received their first frogs in 2003. Since then, they have been able to successfully breed 500 individuals in captivity but will not release them into the wild until the fungal disease is less of a threat. The San Diego zoo also sends money to Panama to keep up the
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The juveniles of this species are amphibious, but have a far smaller range than subadults and adults. Normally, the juveniles are not found more than 2 m from their streams, and recently metamorphosed juveniles are more likely to be found next to the stream pools teeming with tadpoles. Like their
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In early 2006, EVACC exceeded its capacity for housing golden frogs. In order to continue collection efforts, a partnership was formed with the Hotel
Campestre in El Valle whereby rooms 28 and 29 of the resort were loaded with terrariums as a stop-gap measure. This was one of several conservation
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The temperature at which these amphibians keep may be correlated to chytridiomycosis; the fungus is more prevalent in colder conditions. If a cold period occurs, the behavior and immunity of the frogs may change around the same time more spores are released. When these frogs are infected with the
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The
Panamanian golden frog is a national symbol and is considered to be one of the most beautiful frogs in Panama. The skin colour ranges from light yellow-green to bright gold, with some individuals exhibiting black spots on their backs and legs. Females are generally larger than males; females
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Project Golden Frog is a conservation project involving scientific, educational, and zoological institutions in the
Republic of Panama and the United States. The intended outcomes of this project include a greater understanding of the Panamanian golden frog, a coordinated conservation effort by
462:. Wherever water pools in a stream, they are likely to be found, as long as it is connected to moving channels. The tadpoles, however, do not venture into the moving channels. Clinging to surfaces by suction of their flattened bellies, the larvae can be characterized as
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established the El Valle Amphibian Conservation Center (EVACC) in Panama, so the endangered frogs could have protected facilities in their native country. EVACC has become a tourist attraction and the populations of the housed species are watched closely by researchers.
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The Panamanian golden frog began vanishing from its high mountain forests in the late 1990s, prompting a scientific investigation and rescue process that continues today. It was filmed for the last time in the wild in 2006 by the BBC Natural History Unit for the series
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Two significant efforts to save these frogs have been made. The Amphibian Recovery Conservation Coalition, which started in 2004, exported the endangered amphibians to the US, believing it was a better environment for the endangered species. In 2005, the
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typically range from 45 to 63 mm (1.8 to 2.5 in) in length and 4 to 15 g (0.14 to 0.53 oz) in weight, with males between 35 and 48 millimetres (1.4 and 1.9 in) in length and 3 and 12 grams (0.11 and 0.42 oz) in weight.
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the frog's warnings about its toxicity, the foot waving, often accompanied with vocalizations, will continue and increase in frequency and volume. Its toxicity is not a foolproof method of protection, since some animals, such as the
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floods their dermal layers, giving the larvae protection from the sun. When metamorphosing, their golden flecks are replaced with dark green ones. The tadpoles feed on algae and spend 6 to 7 months developing and growing.
608:. Over 300 frogs were kept in the so-called "golden frog hotel" and treated to daily cleansing rinses, 24-hour room service, and exotic lunches of specialty crickets until space could be made available in the EVACC.
570:. The infection is caused by an invasive fungal pathogen that reached El Valle, the home of the Panamanian golden frog, in 2006. Additional factors, such as habitat loss and pollution, may have also played a role.
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Becker, Matthew H.; Richards-Zawacki, Corinne L.; Gratwicke, Brian; Belden, Lisa K. (2014-08-01). "The effect of captivity on the cutaneous bacterial community of the critically endangered Panamanian golden frog
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color which more closely resembles the color of the metamorphic juvenile than the brilliant and sometimes spotted golden color of the adult. The patterning of subadults is significantly darker than the adults.
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Reintroduction efforts from captive assurance colonies are unlikely to be successful without the development of methods to control chytridiomycosis in the wild. Researchers have experimented by using
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Bustamante, Heidi M.; Livo, Lauren J.; Carey, Cynthia (2010). "Effects of temperature and hydric environment on survival of the Panamanian Golden Frog infected with a pathogenic chytrid fungus".
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to control the skin temperature in hopes the fungus would be kept at bay. It seemed to be a protective treatment at the early stage, but the frogs began to die after two weeks as the
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Gagliardo, R.; Griffith, E.; Mendelson, J.; Ross, H.; Zippel, K. (2008). "The principles of rapid response for amphibian conservation, using the programmes in Panama as an example".
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Not only do these frogs face the threat of the fungal disease, but they also are threatened by human development. As trees are cleared for housing and urbanization, the habitat of
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is the causative agent of chytridiomycosis. Other methods of focusing on the phenotypic and genetic concordance to do the conservation have been used. Researchers designated
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261:. Panamanian golden frogs inhabit the streams along the mountainous slopes of the Cordilleran cloud forests of west-central Panama. While the
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The Panamanian golden frog is endemic to Panama, living close to mountain streams on the eastern side of the Tabasará mountain range in the
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that produces a chemical against the infections; however, the skin of Panamanian golden frogs was unsuitable for the bacterium used. The
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1370:
Lindquist, Erik; Thomas Hetherington (1996). "Field studies on visual and acoustic signaling in the "earless" Panamanian golden frog,
1055:"Tests of phenotypic and genetic concordance and their application to the conservation of Panamanian golden frogs (Anura, Bufonidae)"
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550:. The remaining few specimens were taken into captivity and the location of filming was kept secret to protect them from potential
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as separate species, but they are poorly sorted based on physical characteristics, and better sorted based on mitochondrial DNA.
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Populations of amphibians, including the Panamanian golden frog, suffered major declines possibly due to the fungal infection
1992:
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1548:"Thermoregulatory behaviour affects prevalence of chytrid fungal infection in a wild population of Panamanian golden frogs"
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436:; if not, she will attempt to buck him off by arching her spine. Amplexus can last from a few days to a few months, with
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The lifespan of the Panamanian golden frog is 12 years. This toad is unusual in that it communicates by a form of
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Lindquist, Erik; Scott A. Sapoznick; Edgardo J. Griffith Rodriguez; Peter B. Johantgen; Joni M. Criswell (2007).
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The Panamanian golden frog is something of a national symbol, appearing on state lottery tickets and in local
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Fuhrman, F. A.; Fuhrman, G. J.; Mosher, H. S. (29 September 1969). "Toxin from skin of frogs of the genus
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is destroyed. Other threats include encroachment by agriculture, pollution, pet trade, and aquaculture.
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frog, it is difficult to imagine another system of hearing that does not involve an eardrum apparatus.
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Lindquist, Erik; Hetherington, Thomas (1998). "Tadpoles and juveniles of the Panamanian golden frog,
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then grabs the female and hangs on when she crosses his path. If she is receptive, she will tolerate
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423:, waving at rivals and prospective mates, in addition to the sounds more usual among frogs. This
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Woodhams, D. C.; Alford, R. A.; Briggs, C. J.; Johnson, M.; Rollins-Smith, L. A. (2008).
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1506:"Trade-offs influence disease in changing climates: strategies of an amphibian pathogen"
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1890:- Golden Arrow Poison Frog, Golden Frog, Panamanian Golden Frog, Zetek's Golden Frog
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Panama Amphibian Rescue and Conservation Project, 2010 Accessed September 28, 2010
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provinces. Its geographic range previously extended as far east as the town of
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927:(Bufonidae), with information on development of coloration and patterning".
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The Panamanian golden frog has a variety of toxins, including steroidal
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until the functions of the circulatory and respiratory systems cease.
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828:(1933). "Amphibians and reptiles from El Valle de Anton, Panama".
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in western Coclé Province before the onset of the fungal disease
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AmphibiaWeb: Information on amphibian biology and conservation.
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Amphibian Species of the World: an Online Reference. Version 6.0
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1334:(Anura, Bufonidae), with notes on fluorescent pigment tracking"
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in a bid to preserve the species. The alternative common name,
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Occasional Papers of the Boston Society of Natural History
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Despite its common name, the Panamanian golden frog is a
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several orders of magnitude more potent than its analog
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Proceedings of the Royal Society B: Biological Sciences
1684:"Amphibian Rescue and Conservation Project - Panama"
1913:
1330:"Nocturnal position in the Panamanian golden frog,
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1486:. Panama Amphibian Rescue and Conservation Project
1022:
563:conservation effort in the frogs' native country.
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1693:. Amphibians.org. pp. 17–20. Archived from
1053:Richards, Corinne L.; Knowles, L. Lacey (2007).
273:since 2007. Individuals have been collected for
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329:, but is now classified as a separate species.
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780:10.2305/IUCN.UK.2019-3.RLTS.T54563A54341110.en
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755:IUCN SSC Amphibian Specialist Group. (2019).
8:
1824:Happy First Annual National Golden Frog Day!
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1257:
1179:: Differentiation from dendrobatid toxins".
2194:IUCN Red List critically endangered species
1768:"Panama Hotel Is Imperiled Frogs' Lifeboat"
315:, a member of the family Bufonidae. It was
1901:
1745:(First ed.). Henry Holt and Company.
1741:The sixth extinction: an unnatural history
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1168:
605:The Sixth Extinction: An Unnatural History
452:can be divided into four stages: larva or
440:usually taking place in a shallow stream.
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29:
20:
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1349:
1081:
1025:The Amphibians and Reptiles of Costa Rica
970:. Berkeley, California: AmphibiaWeb. 2014
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1766:Roig-Franzia, Manuel (26 October 2006).
1484:"Panama Amphibian Conservation Timeline"
1029:. Chicago: University of Chicago Press.
875:
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1881:Discover Life website: Species ID tool
1876:Discover Life website: Atelopus zeteki
1411:The Case of the Vanishing Golden Frogs
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7:
1897:Panama Travel Guides and Attractions
895:. American Museum of Natural History
360:, has been found to be a blocker of
766:IUCN Red List of Threatened Species
1895:Panamanian Golden Frog Description
1711:Kolbert, Elizabeth (25 May 2009).
1351:10.11606/issn.2316-9079.v6i1p37-44
997:"'Last wave' for wild golden frog"
856:American Museum of Natural History
14:
362:voltage-dependent sodium channels
1649:10.1111/j.1749-4877.2010.00197.x
1546:Richards-Zawacki, C. L. (2009).
1312:10.1111/j.1748-1090.2008.00043.x
1074:10.1111/j.1365-294x.2007.03369.x
680:Decline in amphibian populations
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91:
57:, possibly extinct in the wild
2219:Taxa named by Emmett Reid Dunn
632:Batrachochytrium dendrobatidis
619:Batrachochytrium dendrobatidis
1:
1682:Gratwicke, Brian (May 2012).
1201:10.1126/science.165.3900.1376
2204:Amphibians described in 1933
1886:Global Amphibian Assessment
1811:10.1016/j.biocon.2014.05.029
1737:Kolbert, Elizabeth (2014).
1670:The Conservation Initiative
1273:Endangered Species Bulletin
710:Other common names include
616:Prior to the spread of the
269:, it may in fact have been
244:Cerro Campana stubfoot toad
2235:
1300:International Zoo Yearbook
1232:"The Frogs And The Fungus"
628:Janthinobacterium lividium
556:Janthinobacterium lividium
356:class. One of the latter,
1454:10.1007/s10393-012-0743-0
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88:Scientific classification
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1860:National Zoological Park
1839:"Panamanian golden frog"
1151:"Panamanian Golden Frog"
1119:"Panamanian Golden Toad"
716:Golden arrow poison frog
412:Panamanian golden frogs
2209:Endemic fauna of Panama
1799:Biological Conservation
1713:"The Sixth Extinction?"
1409:Markle, Sandra (2011).
1230:Konstant, Bill (2007).
1123:Encyclopædia Britannica
24:Panamanian golden frog
1564:10.1098/rspb.2009.1656
1376:Journal of Herpetology
999:. BBC. 2 February 2008
500:
416:
308:
246:and other names, is a
234:Panamanian golden frog
221:Atelopus varius zeteki
1266:"Project golden frog"
1264:Poole, Vicky (2008).
773:: e.T54563A54341110.
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411:
303:
285:both commemorate the
275:breeding in captivity
267:Critically Endangered
55:Critically endangered
2214:Amphibians of Panama
1844:Encyclopedia of Life
802:"Appendices | CITES"
691:Ecnomiohyla rabborum
16:Species of amphibian
1867:Species information
1865:Project Golden Frog
1772:The Washington Post
1637:Integrative Zoology
1612:2010Natur.465U.848.
1606:(7300): 848. 2010.
1193:1969Sci...165.1376F
1187:(3900): 1376–1377.
724:Zetek's golden frog
685:Holocene Extinction
598:efforts covered in
528:Liophis epinephalus
448:The development of
427:is thought to have
279:Zetek's golden frog
271:extinct in the wild
40:Conservation status
548:David Attenborough
543:Life in Cold Blood
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498:Vancouver Aquarium
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306:Vancouver Aquarium
281:, and the epithet
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1907:Taxon identifiers
1752:978-0-8050-9299-8
1558:(1681): 519–528.
1525:10.1890/06-1842.1
1420:978-0-7613-5108-5
1068:(15): 3119–3133.
1062:Molecular Ecology
858:. 23 January 2020
600:Elizabeth Kolbert
465:gastromyzophorous
242:), also known as
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398:chytridiomycosis
223:
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1996:
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1983:
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1959:Atelopus_zeteki
1957:
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1945:Atelopus zeteki
1943:
1942:
1937:
1928:
1927:
1922:
1915:Atelopus zeteki
1909:
1888:Atelopus zeteki
1850:Panamanian Frog
1835:
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1795:Atelopus zeteki
1791:
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1440:golden frogs".
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1388:10.2307/1565171
1372:Atelopus zeteki
1369:
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1332:Atelopus zeteki
1327:
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759:Atelopus zeteki
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507:Atelopus varius
487:
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382:
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326:Atelopus varius
317:first described
298:
239:Atelopus zeteki
207:
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197:Atelopus zeteki
195:
182:
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1833:External links
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1717:The New Yorker
1703:
1700:on 2012-08-06.
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1643:(2): 143–153.
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1448:(4): 501–506.
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1382:(3): 347–354.
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1083:2027.42/102716
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1019:Savage, Jay M.
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524:colubrid snake
486:
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358:zetekitoxin AB
343:bufadienolides
338:
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188:Binomial name
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1842:
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1787:
1775:. Retrieved
1771:
1761:
1740:
1732:
1720:. Retrieved
1716:
1706:
1695:the original
1690:
1677:
1665:
1640:
1636:
1630:
1603:
1599:
1590:
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1541:
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1488:. Retrieved
1478:
1445:
1441:
1410:
1404:
1379:
1375:
1371:
1344:(1): 37–44.
1341:
1338:Phyllomedusa
1337:
1331:
1303:
1299:
1293:
1281:. Retrieved
1276:
1272:
1244:. Retrieved
1239:
1235:
1225:
1184:
1180:
1176:
1155:. Retrieved
1126:. Retrieved
1122:
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1061:
1024:
1013:
1001:. Retrieved
972:. Retrieved
967:
961:
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897:. Retrieved
892:
886:
860:. Retrieved
855:
849:
842:
833:
829:
820:
809:. Retrieved
805:
796:
784:. Retrieved
770:
764:
758:
723:
720:Zetek's frog
719:
715:
711:
706:
689:
672:Frogs portal
649:
639:
635:
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535:Conservation
526:
518:
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505:
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479:
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463:
458:
449:
447:
444:Life history
418:
383:
380:Distribution
371:
354:tetrodotoxin
340:
331:
324:
310:
287:entomologist
282:
278:
265:lists it as
243:
238:
237:
233:
231:
220:
219:
196:
194:
178:
177:
165:
18:
2068:iNaturalist
1967:AmphibiaWeb
1939:Wikispecies
1871:Houston Zoo
1805:: 199–206.
1777:30 November
1722:30 November
1003:22 February
889:Dunn, 1933"
852:Dunn, 1933"
712:Golden frog
591:Houston Zoo
438:oviposition
347:guanidinium
296:Description
290:James Zetek
2188:Categories
1490:2009-09-13
1283:14 January
1246:26 October
1157:14 January
1128:26 October
899:28 January
862:23 January
826:Dunn, E.R.
811:2022-01-14
786:11 October
734:References
646:In culture
636:A. various
612:Experiment
425:adaptation
321:subspecies
224:Dunn, 1933
74:Appendix I
1442:EcoHealth
1279:(1): 7–10
1242:(5): 6–12
974:7 October
806:cites.org
652:mythology
640:A. zeteki
579:A. zeteki
519:A. zeteki
459:A. certus
450:A. zeteki
421:semaphore
372:grand mal
366:saxitoxin
350:alkaloids
313:true toad
173:Species:
155:Bufonidae
111:Kingdom:
105:Eukaryota
2199:Atelopus
2156:Species+
2086:10212839
1930:Q2048914
1924:Wikidata
1691:Frog Log
1657:21392332
1582:19864287
1533:18589527
1462:22328095
1236:Americas
1217:41245748
1177:Atelopus
1100:21631940
1092:17651191
1021:(2002).
883:(2016).
836:: 65–79.
658:See also
552:poachers
494:Amplexus
485:Behavior
434:amplexus
370:clonic (
337:Toxicity
213:Synonyms
166:Atelopus
151:Family:
135:Amphibia
125:Chordata
121:Phylum:
115:Animalia
101:Domain:
61:IUCN 3.1
2060:5216695
2047:1039153
1841:at the
1608:Bibcode
1573:2842693
1513:Ecology
1470:8609071
1396:1565171
1209:5807965
1189:Bibcode
1181:Science
941:3893155
470:melanin
454:tadpole
429:evolved
404:Ecology
394:El Copé
352:of the
304:At the
255:endemic
248:species
161:Genus:
141:Order:
131:Class:
76: (
59: (
2168:uBio:
2148:377316
2112:550266
2099:112533
2031:ECOS:
1980:ARKive
1749:
1655:
1600:Nature
1580:
1570:
1531:
1468:
1460:
1417:
1394:
1215:
1207:
1098:
1090:
1033:
939:
722:, and
414:mating
390:Panamá
283:zeteki
259:Panama
206:, 1933
2171:26328
2132:NAS:
2125:54563
2081:IRMNG
2073:21723
2024:5VVKC
2011:81807
1698:(PDF)
1687:(PDF)
1509:(PDF)
1466:S2CID
1392:JSTOR
1269:(PDF)
1213:S2CID
1096:S2CID
1058:(PDF)
937:JSTOR
698:Notes
517:When
386:Coclé
319:as a
145:Anura
78:CITES
72:CITES
2161:7380
2143:NCBI
2120:IUCN
2107:ITIS
2055:GBIF
2006:BOLD
1797:)".
1779:2018
1747:ISBN
1724:2018
1653:PMID
1578:PMID
1529:PMID
1458:PMID
1415:ISBN
1285:2014
1248:2015
1205:PMID
1159:2014
1130:2015
1088:PMID
1031:ISBN
1005:2015
976:2014
901:2016
864:2020
788:2023
771:2019
638:and
388:and
345:and
263:IUCN
252:toad
232:The
204:Dunn
2094:ISC
2042:EoL
2034:779
2019:CoL
1993:ASW
1954:ADW
1807:doi
1803:176
1645:doi
1616:doi
1604:465
1568:PMC
1560:doi
1556:277
1521:doi
1450:doi
1384:doi
1374:".
1346:doi
1308:doi
1197:doi
1185:169
1078:hdl
1070:doi
775:doi
602:'s
546:by
323:of
257:to
250:of
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