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

Holoadeninae Hedges, Duellman, and Heinicke, 2008

Class: Amphibia > Order: Anura > Superfamily: Brachycephaloidea > Family: Craugastoridae > Subfamily: Holoadeninae

154 species

Holoadeninae Hedges, Duellman, and Heinicke, 2008, Zootaxa, 1737: 97. Type genus: Holoaden Miranda-Ribeiro, 1920.

Hypodactylinae Heinicke, Lemmon, Lemmon, McGrath, and Hedges, 2017 "2018", Mol. Phylogenet. Evol., 118: 145. Type genus: Hypodactylus Hedges, Duellman, and Heinicke, 2008.

English Names

None noted.

Distribution

Pacific lowlands of Ecuador and southern Colombia; in the Andes of southern Ecuador, Peru, and Bolivia, and in the Amazon Basin of Colombia, Peru, Bolivia, and adjacent Brazil; Atlantic coastal forests in southeastern Brazil.

Comment

Association of Atopophrynus, Dischidodactylus, Euparkerella, and Geobatrachus was made provisionally with this taxon by Hedges, Duellman, and Heinicke, 2008, Zootaxa, 1737: 1-182, substantially on the basis of geography (p. 97). Padial, Castroviejo-Fisher, and De la Riva, 2009, Mol. Phylogenet. Evol., 52: 911-915, provided molecular evidence for the monophyly of the group. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543-583, in their study of Genbank sequences, redelimited the group, suggested that it might be the sister taxon of Pristimantinae, and excluded taxa not included on the basis of character evidence. Canedo and Haddad, 2012, Mol. Phylogenet. Evol., 65: 610-620, suggested that Strabomantinae may be the sister taxon of Holoadeninae. Padial, Grant, and Frost, 2014, Zootaxa, 3825: 53, transferred Hypodactylus, Lynchius, Oreobates, and Phrynopus into this subfamily, noted that no morphological synapomorphies are apparent although they did discuss relationships uncovered by molecular analysis that had been previously discovered by study of morphology. Venegas, Barboza, De la Riva, and Padial, 2018, Zootaxa, 4446: 501–524, provided a tree of the members of Holoadeninae. Catenazzi, Mamani, Lehr, and von May, 2020, Diversity, 12 (184): 1–17, reported on molecular phylogenetics of the group.

Contained taxa (154 sp.):

"Eleutherodactylus" bilineatus (Bokermann, 1975)

Barycholos Heyer, 1969 (2 sp.)

Bryophryne Hedges, Duellman, and Heinicke, 2008 (11 sp.)

Euparkerella Griffiths, 1959 (5 sp.)

Holoaden Miranda-Ribeiro, 1920 (4 sp.)

Lynchius Hedges, Duellman, and Heinicke, 2008 (7 sp.)

Microkayla De la Riva, Chaparro, Castroviejo-Fisher, and Padial, 2017 (25 sp.)

Niceforonia Goin and Cochran, 1963 (15 sp.)

Noblella Barbour, 1930 (16 sp.)

Oreobates Jiménez de la Espada, 1872 (25 sp.)

Phrynopus Peters, 1873 (35 sp.)

Psychrophrynella Hedges, Duellman, and Heinicke, 2008 (5 sp.)

Qosqophryne Catenazzi, Mamani, Lehr, and von May, 2020 (3 sp.)

External links:

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To search the NIH genetic sequence database, see GenBank
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