Search tool	Example of search string in box	What that means to the database
* = "wildcard". Allows a partial letter string to be entered and implies either no or various extension on this letter string	Hyl*	Will find all uses of "Hyl . . ." anywhere in a record: e.g., Hylarana, Hyla, Hylidae, Hylinae, Hylaedactyla.
* = "wildcard". Allows a partial letter string to be entered and implies either no or various extension on this letter string	*Hyla	Will find all uses of ". . . hyla" anywhere in a record: e.g., Hyla, Hylidae, Plectrohyla, Ptychadena hylaea, Adenomera hylaedactyla
	Hyla	Will find all records that contain stand-alone uses of Hyla: e.g., Hyla, Hyla arenicolor
Strings of words without quotation marks	Lithobates pipiens	Interprets this as "Lithobates OR pipiens" so will find the union of all records that contain either "Lithobates" or "pipiens": e.g., Lithobates omiltemanus, Hylorana pipiens
Strings of word with quotation marks	"Lithobates pipiens"	Interprets this as "Lithobates AND pipiens" so will return all records that have the character string "Lithobates pipiens" anywhere within a record: e.g., all members of the Lithobates pipiens complex.
AND statement. Means that the results of the search must include both words held together by an AND	Lithobates AND pipiens	Same as "Lithobates pipiens"
OR statement. Means that the results of the search will all records that have either of the terms held together by an OR	Lithobates OR "Costa Rica"	Recovers all records that contain the word "Lithobates" or "Costa Rica": 402 records inasmuch as this will include all records for the entire amphibian fauna of Costa Rica
Combining Boolean operators within the Basic Search box (AND, OR, NOT, quotation marks)	Lithobates AND "Costa Rica" NOT forreri	Recovers only records that contain the word Lithobates AND "Costa Rica", but excludes any record that contains the word "forreri" in any part of the record: 9 records
Printable HELP file

Search tool	Example of search string in box	What that means to the database
* = "wildcard". Allows a partial letter string to be entered and implies either no or various extension on this letter string	Hyl*	Will find all uses of "Hyl . . ." anywhere in a record: e.g., Hylarana, Hyla, Hylidae, Hylinae, Hylaedactyla.
* = "wildcard". Allows a partial letter string to be entered and implies either no or various extension on this letter string	*Hyla	Will find all uses of ". . . hyla" anywhere in a record: e.g., Hyla, Hylidae, Plectrohyla, Ptychadena hylaea, Adenomera hylaedactyla
Single word	Hyla	Will find all records that contain stand-alone uses of Hyla: e.g., Hyla, Hyla arenicolor
Strings of words without quotation marks	Lithobates pipiens	Interprets this as "Lithobates OR pipiens" so will find the union of all records that contain either "Lithobates" or "pipiens": e.g., Lithobates omiltemanus, Hylorana pipiens
Strings of word with quotation marks	"Lithobates pipiens"	Interprets this as "Lithobates AND pipiens" so will return all records that have the character string "Lithobates pipiens" anywhere within a record: e.g., all members of the Lithobates pipiens complex.
AND statement. Means that the results of the search must include both words held together by an AND	Lithobates AND pipiens	Same as "Lithobates pipiens"
OR statement. Means that the results of the search will all records that have either of the terms held together by an OR	Lithobates OR "Costa Rica"	Recovers all records that contain the word "Lithobates" or "Costa Rica": 402 records inasmuch as this will include all records for the entire amphibian fauna of Costa Rica
Combining Boolean operators within the Basic Search box (AND, OR, NOT, quotation marks)	Lithobates AND "Costa Rica" NOT forreri	Recovers only records that contain the word Lithobates AND "Costa Rica", but excludes any record that contains the word "forreri" in any part of the record: 9 records
Printable HELP file

Atelopus varius (Lichtenstein and Martens, 1856)

Class: Amphibia > Order: Anura > Family: Bufonidae > Genus: Atelopus > Species: Atelopus varius

Atelopus varius Stannius, 1856, Handb. Zootomie Wiebelthiere, 2: 16. Nomen nudum.

Phrynidium varium Lichtenstein and Martens, 1856, Nomencl. Rept. Amph. Mus. Zool. Berol.: 40. Syntypes: Not designated; according to museum records: ZMB 3377 (2 specimens) and 3378 (2 specimens) and 3379; ZMB 3377 designated neotype by Savage, 1972, Herpetologica, 28: 84, but this number applies to two specimens, according to Lötters, 1996, Neotrop. Toad Genus Atelopus: 53. Female of ZMB3377 (subsequently 3377.1) designated lectotype by Lötters, Böhme, and Günther, 1998, Mitt. Mus. Naturkd. Berlin, Zool., 74: 176. Type locality: "Veragoa"; discussed by Savage, 1972, Herpetologica, 28: 89. Restricted to "the Pacific portion of the Provincia Veraguas, western Panamá" by Lötters, Böhme, and Günther, 1998, Mitt. Mus. Naturkd. Berlin, Zool., 74: 176.

Phrynidium varium var. adspersum Lichtenstein and Martens, 1856, Nomencl. Rept. Amph. Mus. Zool. Berol.: 41. Syntypes: ZMB 3377-78 (4 specimens); ZMB 3377 designated lectotype by Savage, 1972, Herpetologica, 28: 83; but this number applies to two specimens, according to Lötters, 1996, Neotrop. Toad Genus Atelopus: 53. Female of ZMB 3377 (subsequent. 3377.1) designated lectotype by Lötters, Böhme, and Günther, 1998, Mitt. Mus. Naturkd. Berlin, Zool., 74: 176. Type locality: "Veragoa" = Veragua, Panama; discussed by Savage, 1972, Herpetologica, 28: 89. Synonymy by Dunn, 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 394, and Taylor, 1952, Univ. Kansas Sci. Bull., 35: 624.

Phrynidium varium var. maculatum Lichtenstein and Martens, 1856, Nomencl. Rept. Amph. Mus. Zool. Berol.: 40. Holotype: ZMB 3379, according to Savage, 1972, Herpetologica, 28: 83. Type locality: "Veragoa" = Veragua, Panama; discussed by Savage, 1972, Herpetologica, 28: 89. Restricted to "the western portion of the Provincia Chiriquí, western Panamá" by Lötters, Böhme, and Günther, 1998, Mitt. Mus. Naturkd. Berlin, Zool., 74: 176. Synonymy by Dunn, 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 397, and Lötters, Böhme, and Günther, 1998, Mitt. Mus. Naturkd. Berlin, Zool., 74: 176.

Hylaemorphus bibronii Schmidt, 1857, Sitzungsber. Akad. Wiss. Wien, Phys. Math. Naturwiss. Kl., 24: 14. Holotype: KM 1015/1355, according to Savage, 1970, Proc. California Acad. Sci., Ser. 4, 38: 273-288. Type locality: "Neu-Granada"; later restricted by Schmidt, 1858, Denkschr. Akad. Wiss. Wien, Math. Naturwiss. Kl., 14: 256, to "Neu-Granada, unweit Panama, zwischen 2000′ und 3000′ [Polish feet, therefore = 460-760 m, according to Savage, 1970, Proc. California Acad. Sci., Ser. 4, 38: 273-288]". Synonymy by Cochran and Goin, 1970, Bull. U.S. Natl. Mus., 288: 137. See comment by Lötters, 1996, Neotrop. Toad Genus Atelopus: 53, about type locality and status.

Hylaemorphus dumerilii Schmidt, 1857, Sitzungsber. Akad. Wiss. Wien, Phys. Math. Naturwiss. Kl., 24: 14. Holotype: KM 1014/1355, according to Savage, 1970, Proc. California Acad. Sci., Ser. 4, 38: 273-288. Type locality: "Neu-Granada", later restricted by Schmidt, 1858, Denkschr. Akad. Wiss. Wien, Math. Naturwiss. Kl., 14: 256, to "Neu-Granada, Provinz Veragua, in Höhe bis gegen 8000′". Synonymy by Günther, 1859 "1858", Cat. Batr. Sal. Coll. Brit. Mus.: 137; Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 152.

Hylaemorphus plutonius Jan, 1857, Cenni Mus. Civ. Milano: 53. Type(s): MSNM. Type locality: "Brasile" (presumably in error). Nomen nudum attributed to Fitzinger and Tschudi, presumably on the basis of label names. Presumed to be in this synonymy on the basis of similarity of name with Hylaemorphus pluto Schmidt, 1858 (DRF).

Hylaemorphus pluto Schmidt, 1858, Denkschr. Akad. Wiss. Wien, Math. Naturwiss. Kl., 14: 255. Syntypes: NHMW 3875.1-4 (4 specimens), according to Savage, 1972, Herpetologica, 28: 83. Type locality: "Costa Rica". Restricted to "Cartago, Cantón Cartago, Provincia Cartago, Costa Rica" by Savage, 1974, Rev. Biol. Tropical, 22: 84, who also commented on the type locality. Synonymy by Taylor, 1952, Univ. Kansas Sci. Bull., 35: 624.

Phryniscus bibronii — Günther, 1859 "1858", Cat. Batr. Sal. Coll. Brit. Mus.: 155.

Phryniscus varius — Günther, 1859 "1858", Cat. Batr. Sal. Coll. Brit. Mus.: 45.

Phrynidium bibronii — Cope, 1867, J. Acad. Nat. Sci. Philadelphia, Ser. 2, 6: 196.

Atelopus varius — Keferstein, 1867, Nachr. Ges. Wiss. Göttingen, 18: 350; Boulenger, 1894, Ann. Mag. Nat. Hist., Ser. 6, 14: 374-375.

Atelopus varius var. maculatus — Peters, 1873, Monatsber. Preuss. Akad. Wiss. Berlin, 1873: 618.

Phryniscus dumerilii — Brocchi, 1882, Miss. Scient. Mex. Amer. Centr., Rech. Zool., 3(2, livr. 2): 96.

Phryniscus varius — Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 152.

Atelopus bibronii — Boulenger, 1894, Ann. Mag. Nat. Hist., Ser. 6, 14: 374-375; Rivero, 1963, Caribb. J. Sci., 3: 108.

Atelopus varius varius — Dunn, 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 393-397.

Atelopus varius maculatus — Dunn, 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 397.

Atelopus varius loomisi Taylor, 1952, Univ. Kansas Sci. Bull., 35: 625. Holotype: KU 24746, by original designation. Type locality: "Isla Bonita (American Cinchona plantation), approximately 5000 ft. elevation, eastern slope Volcán Poás, [Cantón de Alajuela, Provincia de Alajuela,] Costa Rica". Type locality commented on by Savage, 1974, Rev. Biol. Tropical, 22: 79.

Atelopus varius ambulatorius Taylor, 1952, Univ. Kansas Sci. Bull., 35: 628. Holotype: R.C. Taylor 766, by original designation; now FMNH 178270 according to Marx, 1976, Fieldiana, Zool., 69: 44. Type locality: "Isla Bonita (American Cinchona Plantation), southeastern slope Volcán Poás, elevation approximately 6000 ft.", Cantón de Alajuela, Provincia de Alajuela, Costa Rica. Type locality commented on by Savage, 1974, Rev. Biol. Tropical, 22: 80.

Atelopus loomisi — Taylor, 1955, Univ. Kansas Sci. Bull., 37: 517. Taxonomic change without discussion.

Atelopus varius bibroni — Cochran and Goin, 1970, Bull. U.S. Natl. Mus., 288: 137.

Common Names

Harlequin Frog (Savage, 1972, Herpetologica, 28: 77–94).

Clown Frog (Trade name).

Painted Frog (Ananjeva, Borkin, Darevsky, and Orlov, 1988, Dict. Amph. Rept. Five Languages: 36).

Veragoa Stubfoot Toad (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 40).

Distribution

Premontane and lower montane zones on both Atlantic and Pacific versants of the cordilleras of Costa Rica and western Panama, below 1600 m elevation (records from eastern Panama and Colombia apparently are based on misidentifications; see comment).

Geographic Occurrence

Natural Resident: Costa Rica, Panama

Comment

In the Atelopus ignescens group of Lynch, 1993, Alytes, 11: 77–87. See account by Savage, 1972, Herpetologica, 28: 77–94. Cocroft, McDiarmid, Jaslow, and Ruiz-Carranza, 1990, Copeia, 1990: 638, suggested that Atelopus varius was a species complex, with at least the Panamanian records represented more than one species. Similarly, Lötters, 1996, Neotrop. Toad Genus Atelopus: 53, suggested that Colombian (source of these is Cochran and Goin, 1970, Bull. U.S. Natl. Mus., 288: 118–145, a notorious source of misidentifications and taxonomic misallocations) and eastern Panamanian records are based on misidentifications. See Lötters, Böhme, and Günther, 1998, Mitt. Mus. Naturkd. Berlin, Zool., 74: 173v184, for discussion of nomenclatural history. See account by Savage, 2002, Amph. Rept. Costa Rica: 189–190, who restricted the distribution to Costa Rica and Panama. Richards and Knowles, 2007, Mol. Ecol., 16: 3119–3133, suggested that at least their samples of Atelopus varius from Costa Rica were more closely related to Atelopus senex than they were to Panamanian Atelopus varius. See photograph, map, description of geographic range and habitat, and conservation status in Stuart, Hoffmann, Chanson, Cox, Berridge, Ramani, and Young, 2008, Threatened Amph. World: 177. Velásquez-Álvarez and Kahn, 2005, in Rueda-Almonacid et al. (eds.), Ranas Arlequines: 115, provided a brief account, photograph, and map. Köhler, 2011, Amph. Cent. Am.: 98–102, compared this to the other species of Central America and provided a map and photograph. Perez, Richards-Zawacki, Krohn, Robak, Griffith, Ross, Gratwicke, Ibáñez D., and Voyles, 2014, Amph. Rept. Conserv., 8: 30–35, reported on populations in western Panama persisting where chytrid fungus is present. Barrio-Amorós, Chaves, and Puschendorf, 2021, Reptiles & Amphibians, 28: 1–15, discussed surviving populations in Costa Rica, identified three distinct morphs, and provided a dot map. Estupiñán, Ferrari, Gonçalves, Barbosa, Vallinoto, and Schneider, 2016, ZooKeys, 637: 89–106, suggested on the basis of COI barcodes that cryptic species are likely. Jiménez-Monge, Montoya-Greenheck, Bolaños, and Alvarado, 2019, Amph. Rept. Conserv., 13 (2: e192): 115–125, discussed a seemingly chytrid-free population in La Amistad International Park, Costa Rica. Veselý and Batista, 2021, Zool. Res., Kunming, 42: 272–279, provided a dot map of the distribution, which suggests that records from eastern Panama and adjacent Colombia are based on misidentifications.

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