Atelopus

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Atelopus
Atelopus certus calling male edit.jpg
Male A. certus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Anura
Family: Bufonidae
Genus: Atelopus
Duméril and Bibron, 1841
Species

See text

Synonyms

AteleopusAgassiz, 1846 (unjustified emendation)

Atelopus is a large genus of Bufonidae, commonly known as harlequin frogs or toads, from Central and South America, ranging as far north as Costa Rica and as far south as Bolivia. Atelopus species are small, generally brightly colored, and diurnal. Most species are associated with mid-to-high elevation streams.

Contents

This genus has been greatly affected by amphibian declines, with about 70% of species now considered endangered or extinct. [1] While threatened by habitat loss, pollution, and introduced species, the primary cause of these declines appears to be the chytrid fungus Batrachochytrium dendrobatidis . [2] [3]

Almost 40% of the described species in the genus are considered possibly extinct; this is raised to 45% when data deficient species are added; this number may be even higher, given that the genus contains many undescribed species that could also be extinct, and many of the species considered critically endangered but extant may have gone extinct after the last surveys that detected them, or could go extinct in the future. [4] For example, there are 32 known Atelopus species (including half a dozen undescribed) in Ecuador. One of these is data deficient (its status is unclear), two are endangered and the remaining are critically endangered. [5] Almost half the Ecuador species have not been recorded in a decade or more and are likely extinct. In some species conservationists have established captive colonies as a safeguard. [5] However, of 80 species that had not been seen since the 1950s, 32 have been sighted in the 21st Century, albeit at dangerously low population numbers. [1] Among the Atelopus species that have been rediscovered decades after their last sighting have been A. arsyescue , A. mindoensis , A. bomolochos, A. ignescens , A. balios , A. longirostris , A. subornatus , A. varius , A. carbonerensis and possibly A. guanujo . The mechanism whereby these species survived extinction remains to be discovered. [1] [6]

New Atelopus species are discovered with some regularity, and many new species have been described in the last decade. Among others, a new subspecies, popularly dubbed the purple fluorescent frog, was discovered in 2007 by scientists Paul Ouboter and Jan Mol during a follow-up survey of the Nassau plateau in Suriname. [7] Leeanne Alonso from Conservation International, the organisation that led the expedition, said this frog may be threatened by illegal gold mining. [8] It was described as a new subspecies of Atelopus hoogmoedi (itself considered a subspecies of A. spumarius by some), named A. h. nassaui in 2012. [9] Two new species were also described in 2020: A. manauensis and A. moropukaqumir , both of which are highly threatened by the chytrid fungus and habitat destruction . [10] [11] Another new species, A. frontizero , was described in 2021. [12]

Species

ImageCommon nameBinomial nameConservation status
Andes stubfoot toad Atelopus andinus Rivero, 1968EN IUCN
Angelito stubfoot toad Atelopus angelito Ardila-Robayo and Ruiz-Carranza, 1998CR IUCN (possibly extinct)
Ardila's stubfoot toad Atelopus ardila Coloma, Duellman, Almendariz, Ron, Teran-Valdez, and Guayasamin, 2010CR IUCN (possibly extinct)
Atelopus arsyecue imported from iNaturalist photo 57985228 on 4 August 2020.jpg Starry night harlequin toad Atelopus arsyecue Rueda-Almonacid, 1994CR IUCN
Atelopus arthuri 56814765.jpg Arthur's stubfoot toad Atelopus arthuri Peters, 1973CR IUCN
Atelopus balios.jpg Rio Pescado stubfoot toad Atelopus balios Peters, 1973CR IUCN
Atelopus barbotini cropped.png Purple harlequin toad Atelopus barbotini Lescure, 1981LC IUCN
Atelopus bomolochos.jpg Azuay stubfoot toad Atelopus bomolochos Peters, 1973CR IUCN
Boulenger's stubfoot toad Atelopus boulengeri Peracca, 1904CR IUCN
Atelopus calima Velásquez Trujillo, Castro Herrera, Lötters & Plewnia, 2024
Rio Carauta stubfoot toad Atelopus carauta Ruiz-Carranza & Hernández-Camacho, 1978DD IUCN
Atelopus carbonerensis.jpg Venezuelan yellow frog or La Carbonera stubfoot toad Atelopus carbonerensis Rivero, 1974CR IUCN
Atelopus carrikeri01.jpg Guajira stubfoot toad Atelopus carrikeri Ruthven, 1916EN IUCN
Atelopus certus orange.jpg Darien stubfoot toad or Toad Mountain harlequin frog Atelopus certus Barbour, 1923CR IUCN
Chiriqui harlequin frog - Atelopus chiriquiensis.jpg Chiriqui harlequin frog Atelopus chiriquiensis Shreve, 1936EX IUCN
Atelopus chirripoensis Savage and Bolaños, 2009DD IUCN
Chocó stubfoot toad Atelopus chocoensis Lötters, 1992CR IUCN (possibly extinct)
Atelopus chrysocorallus La Marca, 1996CR IUCN
Atelopus coynei Andreas Kay.jpg Rio Faisanes stubfoot toad Atelopus coynei Miyata, 1980CR IUCN
Atelopus cruciger imported from iNaturalist photo 83335906 on 4 August 2020 (cropped).jpg Veragua stubfoot toad Atelopus cruciger (Lichtenstein & Martens, 1856)CR IUCN
Atelopus dimorphus Lötters, 2003DD IUCN
Huila stubfoot toad Atelopus ebenoides Rivero, 1963 (possibly extinct)CR IUCN (possibly extinct)
Atelopus gracilis (1).jpg Elegant stubfoot toad Atelopus elegans (Boulenger, 1882)EN IUCN
Peruvian harlequin frog Atelopus epikeisthos Lötters, Schulte & Duellman, 2005EN IUCN
Carabaya stubfoot toad Atelopus erythropus Boulenger, 1903CR IUCN (possibly extinct)
Malvasa stubfoot toad Atelopus eusebianus Rivero & Granados-Díaz, 1993CR IUCN (possibly extinct)
Atelopus eusebiodiazi Venegas, Catenazzi, Siu-Ting & Carrillo, 2008CR IUCN (possibly extinct)
Atelopus exiguus.jpg Mazán jambato frog Atelopus exiguus Boettger, 1892EN IUCN
Atelopus famelicus Rivero & Morales, 1995CR IUCN
Forest stubfoot toad Atelopus farci Lynch, 1993CR IUCN (possibly extinct)
Cayenne Stubfoot Toad (Atelopus flavescens) (10672861725).jpg Cayenne stubfoot toad Atelopus flavescens Duméril & Bibron, 1841VU IUCN
Central Coast Stubfoot Toad (Atelopus franciscus) (10596338993).jpg Central Coast stubfoot toad Atelopus franciscus Lescure, 1974LC IUCN
Border harlequin frog Atelopus frontizero Veselý & Batista, 2021
Antado stubfoot toad Atelopus galactogaster Rivero & Serna, 1993DD IUCN
Giant stubfoot toad Atelopus gigas Coloma, Duellman, Almendáriz, Ron, Terán-Valdez, and Guayasamin, 2010CR IUCN (possibly extinct)
2012-02-08 Post-metamorph Atelopus gylphus - Pirre Harlequin frog.jpg Pirri Range harlequin frog Atelopus glyphus Dunn, 1931CR IUCN
Guanujo stubfoot toad Atelopus guanujo Coloma, 2002CR IUCN (possibly extinct)
La Guitarra stubfoot toad Atelopus guitarraensis Osorno-Muñoz, Ardila-Robayo & Ruiz-Carranza, 2001DD IUCN
Morona-Santiago stubfoot toad Atelopus halihelos Peters, 1973CR IUCN (possibly extinct)
Atelopus hoogmoedi 12982604.jpg Hoogmoed harlequin toad Atelopus hoogmoedi Lescure, 1974
Atelopus ignescens 2019.jpg Quito stubfoot toad Atelopus ignescens (Cornalia, 1849)CR IUCN
Atelopus laetissimus.jpg San Lorenzo stubfoot toad Atelopus laetissimus Ruiz-Carranza, Ardila-Robayo & Hernández-Camacho, 1994EN IUCN
Atelopus limosus (5777160370).jpg Limosa harlequin frog Atelopus limosus Ibáñez, Jaramillo & Solís, 1995CR IUCN
Atelopus loettersi De la Riva, Castroviejo-Fisher, Chaparro, Boistel, and Padial, 2011NT IUCN
El Tambo stubfoot toad Atelopus longibrachius Rivero, 1963EN IUCN
Atelopus longirostris male.jpg Longnose stubfoot toad Atelopus longirostris Cope, 1868CR IUCN
Atelopus lozanoi Osorno-Muñoz, Ardila-Robayo & Ruiz-Carranza, 2001 (endangered)EN IUCN
Lynch's stubfoot toad Atelopus lynchi Cannatella, 1981CR IUCN (possibly extinct)
Manaus harlequin frog Atelopus manauensis Jorge, Ferrão & Lima, 2020
Atelopus mandingues Osorno-Muñoz, Ardila-Robayo & Ruiz-Carranza, 2001DD IUCN
Atelopus marinkellei Cochran and Goin, 1970EN IUCN
Mindo stubfoot toad Atelopus mindoensis Peters, 1973CR IUCN
Colombian stubfoot toad Atelopus minutulus Ruiz-Carranza, Hernández-Camacho & Ardila-Robayo, 1988CR IUCN (possibly extinct)
Mittermeier's stubfoot toad Atelopus mittermeieri Acosta-Galvis, Rueda-Almonacid, Velásquez-Álvarez, Sánchez-Pacheco & Peña Prieto, 2006EN IUCN
Hernandez's stubfoot toad Atelopus monohernandezii Ardila-Robayo, Osorno-Muñoz & Ruiz-Carranza, 2002CR IUCN (possibly extinct)
Atelopus moropukaqumir Herrera-Alva, Díaz, Castillo, Rodolfo & Catenazzi, 2020
Atelopus mucubajiensis.jpg Mucubaji stubfoot toad Atelopus mucubajiensis Rivero, 1974CR IUCN
La Arboleda stubfoot toad Atelopus muisca Rueda-Almonacid & Hoyos, 1992CR IUCN
Atelopus nahumae.jpg Atelopus nahumae Ruiz-Carranza, Ardila-Robayo & Hernández-Camacho, 1994EN IUCN
Atelopus nanay.jpg Sad harlequin frog Atelopus nanay Coloma, 2002CR IUCN
Gualecenita stubfoot toad Atelopus nepiozomus Peters, 1973EN IUCN
Niceforo's stubfoot toad Atelopus nicefori Rivero, 1963CR IUCN (possibly extinct)
Nocturnal harlequin toad Atelopus nocturnus Bravo-Valencia and Rivera-Correa, 2011CR IUCN
Atelopus onorei Coloma, Lötters, Duellman & Miranda-Leiva, 2007CR IUCN (possibly extinct)
Atelopus orcesi Coloma, Duellman, Almendáriz, Ron, Terán-Valdez & Guayasamin, 2010CR IUCN (possibly extinct)
Atelopus oxapampae Lehr, Lötters, and Lundberg, 2008EN IUCN
Rednose stubfoot toad Atelopus oxyrhynchus Boulenger, 1903CR IUCN (possibly extinct)
Schmidt's stubfoot toad Atelopus pachydermus (Schmidt, 1857)CR IUCN (possibly extinct)
Atelopus palmatus 67424333.jpg Andersson's stubfoot toad Atelopus palmatus Andersson, 1945CR IUCN
Pastuso harlequin frog Atelopus pastuso Andersson, 1945CR IUCN (possibly extinct)
Atelopus patazensis03.jpg Pataz stubfoot toad Atelopus patazensis Venegas, Catenazzi, Siu-Ting & Carrillo, 2008CR IUCN
San Isidro stubfoot toad Atelopus pedimarmoratus Rivero, 1963CR IUCN (possibly extinct)
Peru stubfoot toad Atelopus peruensis Gray & Cannatella, 1985CR IUCN (possibly extinct)
Atelopus petersi imported from iNaturalist photo 56815472 on 4 August 2020.jpg Peters' stubfoot toad Atelopus petersi Coloma, Lötters, Duellman & Miranda-Leiva, 2007CR IUCN (possibly extinct)
Painted stubfoot toad Atelopus petriruizi Ardila-Robayo, 1999CR IUCN (possibly extinct)
Atelopus pictiventris Kattan, 1986CR IUCN (possibly extinct)
Pinango stubfoot toad Atelopus pinangoi Rivero, 1982CR IUCN (possibly extinct)
Napo stubfoot toad Atelopus planispina Jiménez de la Espada, 1875CR IUCN (possibly extinct)
Podocarpus stubfoot toad Atelopus podocarpus Coloma, Duellman, Almendáriz, Ron, Terán-Valdez, and Guayasamin, 2010CR IUCN
Atelopus pulcher 1.jpg Río Huallaga stubfoot toad Atelopus pulcher Boulenger, 1882VU IUCN
Atelopus pyrodactylus Venegas & Barrio, 2006CR IUCN
Quimbaya toad Atelopus quimbaya Ruiz-Carranza & Osorno-Muñoz, 1994CR IUCN (possibly extinct)
Atelopus reticulatus Lötters, Haas, Schick & Böhme, 2002DD IUCN
Anori stubfoot toad Atelopus sanjosei Rivero & Serna, 1989CR IUCN
Upper Amazon stubfoot toad Atelopus seminiferus Cope, 1874EN IUCN
Pass stubfoot toad Atelopus senex Taylor, 1952EX IUCN
Baldias harlequin toad Atelopus sernai Ruiz-Carranza & Osorno-Muñoz, 1994CR IUCN (possibly extinct)
Camouflaged harlequin toad Atelopus simulatus Ruiz-Carranza & Osorno-Muñoz, 1994CR IUCN (possibly extinct)
Atelopus siranus Lötters &Henzl, 2000DD IUCN
Sonsón stubfoot toad Atelopus sonsonensis Vélez-Rodriguez & Ruiz-Carranza, 1997CR IUCN (possibly extinct)
Cloud forest stubfoot toad Atelopus sorianoi La Marca, 1983CR IUCN (possibly extinct)
Atelopus spumarius.jpg Pebas stubfoot toad Atelopus spumarius Cope, 1871VU IUCN
Atelopus spurrelli.jpg Condoto stubfoot toad Atelopus spurrelli Boulenger, 1914NT IUCN
Bogota stubfoot toad Atelopus subornatus Werner, 1899CR IUCN
Atelopus tamaense.jpg Venezuela stubfoot toad Atelopus tamaensis La Marca, García-Pérez & Renjifo, 1990CR IUCN
Atelopus tricolor dorsal.jpg Three-colored stubfoot toad Atelopus tricolor Boulenger, 1902CR IUCN
Atelopus varius 1.jpg Veragoa stubfoot toad Atelopus varius (Lichtenstein & Martens, 1856)CR IUCN
Maracay harlequin frog Atelopus vogli Müller, 1934EX IUCN
Walker's stubfoot toad Atelopus walkeri Rivero, 1963DD IUCN
Atelopus zeteki01b.jpg Panamanian golden frog Atelopus zeteki Dunn, 1933CR IUCN (possibly extinct in the wild)

Related Research Articles

<i>Atelopus longirostris</i> Species of amphibian

Atelopus longirostris is a species of harlequin frog, a member of the family of true toads (Bufonidae). It has been recorded only in northern Ecuador. Records from Colombia probably represent different species. Once listed as extinct by the IUCN, it was rediscovered in 2016 after more than two decade with no sightings.

<span class="mw-page-title-main">Panamanian golden frog</span> Species of amphibian

The Panamanian golden frog, also known as Cerro Campana stubfoot toad and other names, is a species of toad endemic to Panama. Panamanian golden frogs inhabit the streams along the mountainous slopes of the Cordilleran cloud forests of west-central Panama. While the IUCN lists it as Critically Endangered, it may in fact have been extinct in the wild since 2007. Individuals have been collected for breeding in captivity in a bid to preserve the species. The alternative common name, Zetek's golden frog, and the epithet zeteki both commemorate the entomologist James Zetek.

<i>Atelopus varius</i> Species of amphibian

Atelopus varius, the Costa Rican variable harlequin toad or clown frog, is a small Neotropical true toad from the family Bufonidae. Once ranging from Costa Rica to Panama, A. varius is now listed as critically endangered and has been reduced to a single remnant population near Quepos, Costa Rica, and has only relict populations in western Panama. Recent variation in air temperature, precipitation, stream flow patterns, and the subsequent spread of a pathogenic chytrid fungus linked to global climate change have been the leading cause of decline for A. varius. A. zeteki has been considered a subspecies of A. varius, but is now generally considered a separate species.

<span class="mw-page-title-main">Starry night toad</span> Species of amphibian

The starry night harlequin toad is a species of toad in the family Bufonidae endemic to the Sierra Nevada de Santa Marta, Colombia. Its natural habitats are sub-Andean and Andean forests, sub-páramo and páramo at 2,000–3,500 m (6,600–11,500 ft) above sea level. It is named after its unique coloration, being largely black with white spots.

<i>Atelopus balios</i> Species of amphibian

Atelopus balios, the Rio Pescado stubfoot toad, is a species of toad in the family Bufonidae. It is endemic to southwestern Ecuador, with records from Pacific lowlands in Azuay, Cañar, and Guayas Provinces. It is a rare species that was already suspected to be extinct, but a single specimen was discovered in 2011 by a team from Conservation International during a hunt for missing amphibians. The decline in amphibian populations is well documented. The Atelopus balios is Critically Endangered as a result of the widespread amphibian Chytridiomycosis fungus that has decimated other amphibian populations. There are only 10 known findings of the tadpole, Atelopus balios.

<span class="mw-page-title-main">Guajira stubfoot toad</span> Species of amphibian

The Guajira stubfoot toad or Carrikeri harlequin frog is a species of toad in the family Bufonidae. It is about 5 cm (2.0 in) long and typically black, though some populations have orange coloration. This species is endemic to the Sierra Nevada de Santa Marta mountain range of northern Colombia. It is critically endangered because of the chytrid fungus, Batrachochytrium dendrobatidis, and habitat destruction due to agriculture. The species had not been seen from 1994 until it was rediscovered in early 2008.

<i>Atelopus cruciger</i> Species of amphibian

Atelopus cruciger, also known as the Veragua stubfoot toad or Rancho Grande harlequin frog, is a species of toad in the family Bufonidae. It is endemic to Venezuela and is known from the central Venezuelan Coastal Range. The species was already suspected to be extinct because, despite considerable effort, none had been found since 1986. However, in 2003, a small population was found, with few other locations discovered later. It is mainly threatened by chytridiomycosis. It is locally called sapito rayado.

<i>Atelopus glyphus</i> Species of amphibian

Atelopus glyphus, the Pirri harlequin frog or Pirri Range stubfoot toad, is a species of toad in the family Bufonidae found in Colombia and Panama within the Northwestern Andean montane forests. Its natural habitats are subtropical or tropical moist montane forests and rivers.

<i>Atelopus ignescens</i> Species of amphibian

Atelopus ignescens, the Jambato toad or Quito stubfoot toad or Jambato harlequin frog, is a species of toad in the family Bufonidae. It is endemic to the northern Andes of Ecuador. This once abundant species was believed to be extinct until its rediscovery in 2016. The specific name ignescens means "to catch fire," presumably in reference to the orange ventral color of this species.

<i>Atelopus limosus</i> Species of amphibian

Atelopus limosus, the limosa harlequin frog is an endangered species of toad in the family Bufonidae endemic to Panama. Its natural habitats are stream banks in tropical moist lowland forests and rivers of the Chagres watershed in central Panama.

Atelopus lynchi, also known as Lynch's stubfoot toad or Lynch's harlequin frog, is a species of toad in the family Bufonidae. It occurs in northern Ecuador (Carchi) and in southern Colombia. It occurs on the Pacific slope of the Cordillera Occidental, as the westernmost ranges of the Andes are known in both Colombia and in Ecuador. Prior to its description, it was confused with Atelopus longirostris.

Atelopus planispina is a species of toad in the family Bufonidae. It is endemic to the eastern slopes of the Andes of Ecuador. According to the IUCN SSC Amphibian Specialist Group, it is restricted to the Napo Province, although other sources suggest somewhat wider range. It has not been seen since 1985 and might already be extinct. Common names Planispina's harlequin frog, Napo stubfoot toad, and flat-spined atelopus have been coined for it.

Atelopus vogli is an extinct species of harlequin frog in the family Bufonidae. It was endemic to Venezuela. It is known from collections in two localities: its type locality, Las Peñas near Hacienda la Trinidad in Aragua, and Montalbán in Carabobo. It was first described as subspecies of Atelopus cruciger. The specific name vogli honours Cornelius Vogl, German priest who was a missionary in Venezuela in 1925–1959. Common name Vogl's harlequin toad has been coined for it.

<span class="mw-page-title-main">Golden toad</span> Extinct species of toad that was endemic to Costa Rica

The golden toad is an extinct species of true toad that was once abundant in a small, high-altitude region of about 4 square kilometres (1.5 sq mi) in an area north of the city of Monteverde, Costa Rica. It was endemic to elfin cloud forest. Also called the Monte Verde toad, Alajuela toad and orange toad, it is commonly considered the "poster child" for the amphibian decline crisis. This toad was first described in 1966 by herpetologist Jay Savage. The last sighting of a single male golden toad was on 15 May 1989, and it has since been classified as extinct by the International Union for Conservation of Nature (IUCN).

<i>Atelopus patazensis</i> Species of amphibian

Atelopus patazensis is a species of toads in the family Bufonidae. It is endemic to Peru and only known from its type locality in Quebrada Los Alisos, near Pataz in the La Libertad Region. There is, however, an unverified observation from Llacuabamba, about 40 km south of Pataz.

Atelopus onorei is a small species of bright yellow and green toads in the family Bufonidae. It is endemic to Ecuador and is only known from the vicinity of its type locality on the western slope of the Cordillera Occidental in the Azuay Province. It has not been seen since 1990 and may now be extinct, but it is possible that overlooked populations remain.

Atelopus pastuso, commonly known as the Pastuso Harlequin Frog, is a species of frog in the family Bufonidae. It has not been seen since 1993, and is possibly extinct.

Atelopus podocarpus is a species of frog in the family Bufonidae. It has not been seen since 1994, and is believed to be possibly extinct.

Atelopus gigas, the Giant Stubfoot Toad, is a species of frog in the family Bufonidae. It has not been seen since 1970 and is considered possibly extinct.

Atelopus orcesi is a species of frog in the family Bufonidae. It has not been seen since 1988 and is possibly extinct.

References

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  2. Lötters, Kielgast, Bielby, Schmidtlein, Bosch, Veith, Walker, Fisher, Rödder (2009). The Link Between Rapid Enigmatic Amphibian Decline and the Globally Emerging Chytrid Fungus. EcoHealth 6(3): 358-372
  3. Stuart, Hoffmann, Chanson, Cox, Berridge, Ramani and Young, editors (2008). Threatened Amphibians of the World. Pp. 100, 160-178. ISBN   978-84-96553-41-5
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  5. 1 2 Tapia, Elicio Eladio; Coloma, Luis Aurelio; Pazmiño-Otamendi, Gustavo; Peñafiel, Nicolás (2017). "Rediscovery of the nearly extinct longnose harlequin frog Atelopus longirostris (Bufonidae) in Junín, Imbabura, Ecuador". Neotropical Biodiversity. 3: 157–167. doi: 10.1080/23766808.2017.1327000 . S2CID   89865746.
  6. "A tale of two frogs: The tough uphill battle for rediscovered species". Mongabay Environmental News. 2024-06-04. Retrieved 2024-06-18.
  7. "MSN".
  8. Zabarenko, Deborah (2007-06-04). "Purple frog among 24 new species found in Suriname". Reuters . Retrieved 2007-05-06.
  9. Amphibians of Suriname, Paul E. Ouboter and Rawien Jairam, Brill 2012, ISBN   978-90-04-21075-2
  10. Herrera-Alva, Valia; Díaz, Vladimir; Castillo, Ernesto; Rodolfo, César; Catenazzi, Alessandro (2020-09-23). "A new species of Atelopus (Anura: Bufonidae) from southern Peru". Zootaxa. 4853 (3): 404–420. doi:10.11646/zootaxa.4853.3.4. ISSN   1175-5334. PMID   33056367. S2CID   222838469.
  11. Jorge, Rafael F.; Ferrão, Miquéias; Lima, Albertina P. (August 2020). "Out of Bound: A New Threatened Harlequin Toad (Bufonidae, Atelopus) from the Outer Borders of the Guiana Shield in Central Amazonia Described through Integrative Taxonomy". Diversity. 12 (8): 310. doi: 10.3390/d12080310 .
  12. Veselý, Milan; Batista, Abel (2021). "A new species of Atelopus (Amphibia: Bufonidae) from eastern Panama". Zoological Research. 42 (3): 272–279. doi: 10.24272/j.issn.2095-8137.2020.319 . ISSN   2095-8137. PMC   8175953 . PMID   33797210.