''Eleutherodactylus'' is a genus of frogs in the family Eleutherodactylidae.Hedges, S. B., W. E. Duellman, and M. P. Heinicke . 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa 1737: 1-182. Many of the 200 species of the genus are commonly known as "rain frogs" or "robber frogs", due to their sharp, high-pitched, insect-like calls. The best-known species is the

common coquí The common coquí or coquí (''Eleutherodactylus coqui'') is a species of frog endemic to Puerto Rico belonging to the family Eleutherodactylidae. The species is named for the loud call the males make at night. This sound serves two purposes. "CO ...

(''E. coqui''), which is both a national symbol of Puerto Rico and a notorious

invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...

in Hawaii. Two ''Eleutherodactylus'' species, '' E. limbatus'' and '' E. iberia'', are among the smallest known frogs, measuring only 8.5 mm in length (only slightly larger than '' Paedophryne amauensis'', which measures around 7.7 mm).

Etymology

The name "Eleutherodactylus" is derived from the Greek words for ‘free-toed’, composed of the Ancient Greek ' (, ‘free, unbound’) and ' (, ‘finger, toe’). Most species are small, slender, and cryptically colored, with three to five free toes. A few, such as the

web-footed coquí The web-footed coqui, stream coqui, Puerto Rican stream frog or Karl's robber frog (''Eleutherodactylus karlschmidti''), is a possibly extinct Puerto Rican frog species in the family Eleutherodactylidae. It was first described by Chapman Grant ...

(''E. karlschmidti'') of Puerto Rico, do have completely webbed feet.

Distribution and habitat

Species of ''Eleutherodactylus'' are found throughout the Neotropics, including the southern United States, Mexico, Central America, and the Caribbean. Additionally, the common coquí (''E. coqui'') has been introduced to several islands in the Hawaiian archipelago, as well as elsewhere in the Pacific. They can be terrestrial, arboreal, or aquatic, typically living in forests or riparian areas, and feeding primarily upon arthropods. Many ''Eleutherodactylus'' species have highly restricted ranges and are found on only one island or in one or a few localities. Even some of these restricted species can occur at very high densities.

Reproduction and development

All species of ''Eleutherodactylus'' are characterized by direct development, in which eggs hatch directly into small frogs, completely bypassing the tadpole stage. This adaptation may be largely responsible for their ecological and evolutionary success. Most species are characterized by parental behaviors, such as egg-guarding by either the male or female parent. In some cases, even young froglets are attended by parents. Another extinct Puerto Rican species, the

golden coquí The golden coquí (''Eleutherodactylus jasperi'') is a rare species of frog endemic to Puerto Rico. General description Golden coquís are roughly in snout-vent length and are olive-gold to yellow-gold without pattern. The juveniles resemble ad ...

(''E. jasperi''), gave birth to live young. Many species (for example, Cook's robber frog, ''E. cooki''), also of Puerto Rico, exhibit sexual dimorphism in size and color. Study on ''Eleutherodactylus'' and '' Lithobates'' amphibians shows that number of offsprings instead of body size may help to find which species require conservation from being extinct.

Phylogenetics

The basis of forming this genus has been morphological, but sequence comparisons of protein-encoding DNA,

mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is use ...

DNA, and ribosomal

RNA Ribonucleic acid (RNA) is a polymeric molecule essential in various biological roles in coding, decoding, regulation and expression of genes. RNA and deoxyribonucleic acid ( DNA) are nucleic acids. Along with lipids, proteins, and carbohydra ...

have shown geographic range is a much more consistent predictor of cladistics for this group of frogs. The climbing habits of many species have

evolved independently Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...

. All true members of the genus have been clustered into

subgenera In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...

, but many less-related species require more genetic data before they are to be officially classified elsewhere. The theory that the eleutherodactyline colonization of Central America and the

Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean Se ...

from their origins in South America occurred during the Cretaceous has fallen out of favor. The fossil record, combined with molecular clock analyses, indicate the subgenera were probably founded by small groups of individuals by flotsam dispersal during the Eocene or

Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...

epochs. Land bridges would have been limited to facilitating dispersal between West Indian islands, however, the Oligocene division of

Hispaniola Hispaniola (, also ; es, La Española; Latin and french: Hispaniola; ht, Ispayola; tnq, Ayiti or Quisqueya) is an island in the Caribbean that is part of the Greater Antilles. Hispaniola is the most populous island in the West Indies, and th ...

and Cuba resulted in further speciation. The distribution of the subgenus '' Syrrhopus'' is most likely due to a secondary dispersal to Central America from the Greater Antilles during the Miocene. The formation of the

Panama Isthmus The Isthmus of Panama ( es, Istmo de Panamá), also historically known as the Isthmus of Darien (), is the narrow strip of land that lies between the Caribbean Sea and the Pacific Ocean, linking North and South America. It contains the country ...

during the Pliocene has caused some intercontinental distribution among the clades, although only 20 "South American frogs" have ever made it northwards after the original colonization. ''

Sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

'', however, it should exclude clades with distributions south of the Panama Canal.

Fossil record

Frog remains referred to ''Eleutherodactylus'' have been reported from

Dominican amber Dominican amber is amber from the Dominican Republic derived from resin of the extinct tree ''Hymenaea protera''. Dominican amber differentiates itself from Baltic amber by being nearly always transparent, and it has a higher number of fossil incl ...

Species

The following species are recognised in the genus ''Eleutherodactylus'': * '' Eleutherodactylus aporostegus'' Schwartz, 1965 * '' Eleutherodactylus bothroboans'' Schwartz, 1965 * ''

Eleutherodactylus campi ''Eleutherodactylus cystignathoides'', also known as the Rio Grande chirping frog, Mexican chirping frog, or lowland chirping frog, is a small eleutherodactylid frog. It is found from the southern United States in Texas, and in the northeast ...

'' (Stejneger, 1915) * ''

Eleutherodactylus counouspeus ''Eleutherodactylus counouspeus'', also known as Counou robber frog and yellow cave frog, is a species of frog in the family Eleutherodactylidae. It is endemic to the Massif de la Hotte, southwestern Haiti. The specific name ''counouspeus'' refer ...

'' Schwartz, 1964 * ''

Eleutherodactylus diplasius ''Eleutherodactylus diplasius'', the patternless whistling frog, is a species of frog in the family Eleutherodactylidae. A native of Haiti, it can be found in the Tiburon Peninsula northwest, to the Massif de la Hotte. It has an estimated polygon ...

'' Schwartz, 1973 * ''

Eleutherodactylus erythroproctus ''Eleutherodactylus'' is a genus of frogs in the family Eleutherodactylidae.Hedges, S. B., W. E. Duellman, and M. P. Heinicke . 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and ...

'' Schwartz, 1960 * '' Eleutherodactylus feichtingeri'' Díaz, Hedges, and Schmid, 2012 * ''

Eleutherodactylus limbensis ''Eleutherodactylus'' is a genus of frogs in the family Eleutherodactylidae.Hedges, S. B., W. E. Duellman, and M. P. Heinicke . 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and co ...

'' Lynn, 1958 * '' Eleutherodactylus melatrigonum'' Schwartz, 1966 * '' Eleutherodactylus notidodes'' Schwartz, 1966 * ''

Eleutherodactylus olibrus ''Eleutherodactylus'' is a genus of frogs in the family Eleutherodactylidae.Hedges, S. B., W. E. Duellman, and M. P. Heinicke . 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and ...

'' Schwartz, 1958 * ''

Eleutherodactylus orarius ''Eleutherodactylus'' is a genus of frogs in the family Eleutherodactylidae.Hedges, S. B., W. E. Duellman, and M. P. Heinicke . 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and ...

'' (Dixon, 1957) * ''

Eleutherodactylus paralius ''Eleutherodactylus'' is a genus of frogs in the family Eleutherodactylidae.Hedges, S. B., W. E. Duellman, and M. P. Heinicke . 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and co ...

'' Schwartz, 1976 * '' Eleutherodactylus rucillensis'' Cochran, 1939 * ''

Eleutherodactylus sommeri ''Eleutherodactylus'' is a genus of frogs in the family Eleutherodactylidae.Hedges, S. B., W. E. Duellman, and M. P. Heinicke . 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and ...

'' Schwartz, 1977 * '' Eleutherodactylus staurometopon'' Schwartz, 1960 * '' Eleutherodactylus tychathrous'' Schwartz, 1965

West Indian (subgenus '' Eleutherodactylus'')

* '' E. (E.) abbotti'' Cochran, 1923 * '' E. (E.) amplinympha'' Kaiser, Green & Schmid, 1994 * '' E. (E.) antillensis'' Reinhardt & Lütken, 1863 * '' E. (E.) audanti'' Cochran, 1934 * '' E. (E.) auriculatoides'' Noble, 1923 * '' E. (E.) auriculatus'' Cope, 1862 * '' E. (E.) barlagnei'' Lynch, 1965 * '' E. (E.) bartonsmithi'' Schwartz, 1960 * '' E. (E.) brittoni'' Schmidt, 1920 * '' E. (E.) cochranae'' Grant, 1932 * '' E. (E.) cooki'' Grant, 1932 * '' E. (E.) coqui'' Thomas, 1966 * '' E. (E.) eileenae'' Dunn, 1926 * '' E. (E.) eneidae'' Rivero, 1959 * '' E. (E.) flavescens'' Noble, 1923 * '' E. (E.) fowleri'' Schwartz, 1973 * '' E. (E.) glamyrus'' Estrada & Hedges, 1997 * '' E. (E.) gryllus'' Schmidt, 1920 * '' E. (E.) guantanamera'' Hedges, Estrada & Thomas, 1992 * '' E. (E.) haitianus'' Barbour, 1942 * '' E. (E.) hedricki'' Rivero, 1963 * '' E. (E.) ionthus'' Schwartz, 1960 * '' E. (E.) jasperi'' Drewry & Jones, 1976 * '' E. (E.) johnstonei'' Barbour, 1914 * '' E. (E.) juanariveroi'' Ríos-López & Thomas, 2007 * '' E. (E.) karlschmidti'' Grant, 1931 * '' E. (E.) lamprotes'' Schwartz, 1973 * '' E. (E.) leberi'' Schwartz, 1965 * '' E. (E.) locustus'' Schmidt, 1920 * '' E. (E.) mariposa'' Hedges, Estrada & Thomas, 1992 * '' E. (E.) martinicensis'' Tschudi, 1838 * '' E. (E.) melacara'' Hedges, Estrada & Thomas, 1992 * '' E. (E.) minutus'' Noble, 1923 * '' E. (E.) montanus'' Schmidt, 1919 * '' E. (E.) parabates'' Schwartz, 1964 * '' E. (E.) patriciae'' Schwartz, 1965 * '' E. (E.) pinchoni'' Schwartz, 1967 * '' E. (E.) pituinus'' Schwartz, 1965 * '' E. (E.) poolei'' Cochran, 1938 * '' E. (E.) portoricensis'' Schmidt, 1927 * '' E. (E.) principalis'' Estrada & Hedges, 1997 * '' E. (E.) richmondi'' Stejneger, 1904 * '' E. (E.) ronaldi'' Schwartz, 1960 * '' E. (E.) schwartzi'' Thomas, 1966 * '' E. (E.) unicolor'' Stejneger, 1904 * ''E. (E.) varians'' Gundlach & Peters, 1864 * '' E. (E.) wetmorei'' Cochran, 1932 * '' E. (E.) wightmanae'' Schmidt, 1920

West Indian (subgenus '' Euhyas'')

* '' E. (Eu.) acmonis'' Schwartz, 1960 * '' E. (Eu.) adelus'' Diaz, Cadiz & Hedges, 2003 * '' E. (Eu.) albipes'' Barbour & Shreve, 1937 * '' E. (Eu.) alcoae'' Schwartz, 1971 * '' E. (Eu.) alticola'' Lynn, 1937 * '' E. (Eu.) amadeus'' Hedges, Thomas & Franz, 1987 * '' E. (Eu.) andrewsi'' Lynn, 1937 * '' E. (Eu.) apostates'' Schwartz, 1973 * '' E. (Eu.) armstrongi'' Noble & Hassler, 1933 * '' E. (Eu.) atknisi'' Dunn, 1925 * '' E. (Eu.) bakeri'' Cochran, 1935 * '' E. (Eu.) beguei'' Díaz and Hedges, 2015 * '' E. (Eu.) blairhedgesi'' Estrada, Díaz & Rodriguez, 1998 * '' E. (Eu.) bresslerae'' Schwartz, 1960 * '' E. (Eu.) brevirostris'' Shreve, 1936 * '' E. (Eu.) caribe'' Hedges & Thomas, 1992 * '' E. (Eu.) casparii'' Dunn, 1926 * '' E. (Eu.) cavernicola'' Lynn, 1954 * '' E. (Eu.) corona'' Hedges & Thomas, 1992 * '' E. (Eu.) cubanus'' Barbour, 1942 * '' E. (Eu.) cundalli'' Dunn, 1926 * '' E. (Eu.) cuneatus'' Cope, 1862 * '' E. (Eu.) darlingtoni'' Cochran, 1935 * '' E. (Eu.) dimidiatus'' Cope, 1862 * '' E. (Eu.) dolomedes'' Hedges & Thomas, 1992 * '' E. (Eu.) emiliae'' Dunn, 1926 * '' E. (Eu.) etheridgei'' Schwartz, 1958 * '' E. (Eu.) eunaster'' Schwartz, 1973 * '' E. (Eu.) furcyensis'' Shreve & Williams, 1963 * '' E. (Eu.) fuscus'' Lynn & Dent, 1943 * '' E. (Eu.) glandulifer'' Cochran, 1935 * '' E. (Eu.) glanduliferoides'' Shreve, 1936 * '' E. (Eu.) glaphycompus'' Schwartz, 1973 * '' E. (Eu.) glaucoreius'' Schwartz & Fowler, 1973 * '' E. (Eu.) goini'' Schwartz, 1960 * '' E. (Eu.) gossei'' Dunn, 1926 * '' E. (Eu.) grabhami'' Dunn, 1926 * '' E. (Eu.) grahami'' Schwartz, 1979 * '' E. (Eu.) greyi'' Dunn, 1926 * '' E. (Eu.) griphus'' Crombie, 1986 * '' E. (Eu.) guanahacabibes'' Estrada & Rodriguez, 1985 * '' E. (Eu.) gundlachi'' Schmidt, 1920 * '' E. (Eu.) heminota'' Shreve & Williams, 1963 * '' E. (Eu.) iberia'' Estrada & Hedges, 1996 * '' E. (Eu.) intermedius'' Barbour & Shreve, 1937 * '' E. (Eu.) jamaicensis'' Barbour, 1910 * '' E. (Eu.) jaumei'' Estrada & Alonso, 1997 * '' E. (Eu.) jugans'' Cochran, 1937 * '' E. (Eu.) junori'' Dunn, 1926 * '' E. (Eu.) klinikowskii'' Schwartz, 1959 * '' E. (Eu.) lentus'' Cope, 1862 * '' E. (Eu.) leoncei'' Shreve & Williams, 1963 * '' E. (Eu.) limbatus'' Cope, 1862 * '' E. (Eu.) lucioi'' Schwartz, 1980 * '' E. (Eu.) luteolus'' Gosse, 1851 * '' E. (Eu.) maestrensis'' Díaz, Cádiz & Navarro, 2005 * '' E. (Eu.) michaelschmidi'' Díaz, Cádiz & Navarro, 2007 * '' E. (Eu.) monensis'' Meerwarth, 1901 * '' E. (Eu.) nubicola'' Dunn, 1926 * '' E. (Eu.) orcutti'' Dunn, 1928 * '' E. (Eu.) orientalis'' Barbour & Shreve, 1937 * '' E. (Eu.) oxyrhyncus'' Duméril & Bibron, 1841 * '' E. (Eu.) pantoni'' Dunn, 1926 * '' E. (Eu.) paulsoni'' Schwartz, 1964 * '' E. (Eu.) pentasyringos'' Schwartz & Fowler, 1973 * '' E. (Eu.) pezopetrus'' Schwartz, 1960 * '' E. (Eu.) pictissimus'' Cochran, 1935 * '' E. (Eu.) pinarensis'' Dunn, 1926 * '' E. (Eu.) planirostris'' Cope, 1862 * '' E. (Eu.) probolaeus'' Schwartz, 1965 * '' E. (Eu.) rhodesi'' Schwartz, 1980 * '' E. (Eu.) ricordii'' Duméril & Bibron, 1841 * '' E. (Eu.) riparius'' Estrada & Hedges, 1998 * '' E. (Eu.) rivularis'' Diaz, Estrada & Hedges, 2001 * '' E. (Eu.) rogersi'' Goin, 1955 * '' E. (Eu.) rufifemroralis'' Noble & Hassler, 1933 * '' E. (Eu.) schmidti'' Noble, 1923 * '' E. (Eu.) sciagraphus'' Schwartz, 1973 * '' E. (Eu.) semipalmatus'' Shreve, 1936 * '' E. (Eu.) simulans'' Diaz & Fong, 2001 * '' E. (Eu.) sisyphodemus'' Crombie, 1977 * '' E. (Eu.) tetajulia'' Estrada & Hedges, 1996 * '' E. (Eu.) thomasi'' Schwartz, 1959 * '' E. (Eu.) thorectes'' Hedges, 1988 * '' E. (Eu.) toa'' Estrada & Hedges, 1991 * '' E. (Eu.) tonyi'' Estrada & Hedges, 1997 * '' E. (Eu.) turquinensis'' Barbour & Shreve, 1937 * '' E. (Eu.) varleyi'' Dunn, 1925 * '' E. (Eu.) ventrilineatus'' Shreve, 1936 * '' E. (Eu.) warreni'' Schwartz, 1976 * '' E. (Eu.) weinlandi'' Barbour, 1914 * '' E. (Eu.) zugi'' Schwartz, 1958

Hispaniola Hispaniola (, also ; es, La Española; Latin and french: Hispaniola; ht, Ispayola; tnq, Ayiti or Quisqueya) is an island in the Caribbean that is part of the Greater Antilles. Hispaniola is the most populous island in the West Indies, and th ...
n (subgenus '' Pelorius'')

* '' E. (P.) cattus'' Rodriguez, Dugo-Cota, Montero-Mendieta, Gonzalez-Voyer, Alonso Bosch, Vences, and Vilà, 2017 * '' E. (P.) chlorophenax'' Schwartz, 1976 * '' E. (P.) geitonos'' Díaz, Incháustegui, Marte, Köhler, Cádiz, and Rodríguez, 2018 * '' E. (P.) hypostenor'' Schwartz, 1965 * '' E. (P.) inoptatus'' Barbour, 1914 * '' E. (P.) ligiae'' Incháustegui, Díaz, and Marte, 2015 * '' E. (P.) neiba'' Incháustegui, Díaz, and Marte, 2015 * '' E. (P.) nortoni'' Schwartz, 1976 * '' E. (P.) parapelates'' Hedges & Thomas, 1987 * '' E. (P.) ruthae'' Noble, 1923

North/ Central American and Cuban (subgenus '' Syrrhopus'')

* '' E. (S.) albolabris'' Lynch & Lescure, 1980 * '' E. (S.) angustidigitorum'' Taylor, 1940 * ''

Eleutherodactylus colimotl ''Eleutherodactylus'' is a genus of frogs in the family Eleutherodactylidae.Hedges, S. B., W. E. Duellman, and M. P. Heinicke . 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and ...

'' Grünwald, Reyes-Velasco, Franz-Chávez, Morales-Flores, Ahumada-Carrillo, Jones, and Boissinot, 2018 * '' E. (S.) cystingnathoides'' Cope, 1877 * '' E. (S.) dennisi'' Lynch, 1970 * '' E. (S.) dilatus'' Davis & Dixon, 1955 * '' E. (S.) erendirae'' Grünwald, Reyes-Velasco, Franz-Chávez, Morales-Flores, Ahumada-Carrillo, Jones, and Boissinot, 2018 * '' E. (S.) floresvillelai'' Grünwald, Reyes-Velasco, Franz-Chávez, Morales-Flores, Ahumada-Carrillo, Jones, and Boissinot, 2018 * '' E. (S.) grandis'' Dixon, 1957 * '' E. (S.) grunwaldi'' Reyes-Velasco, Ahumada-Carrillo, Burkhardt, and Devitt, 2015 * '' E. (S.) guttilatus'' Cope, 1879 * '' E. (S.) interorbitalis'' Langebartel & Shannon, 1956 * '' E. (S.) jaliscoensis'' Grünwald, Reyes-Velasco, Franz-Chávez, Morales-Flores, Ahumada-Carrillo, Jones, and Boissinot, 2018 * '' E. (S.) leprus'' Cope, 1879 * '' E. (S.) longipes'' Baird, 1859 * '' E. (S.) manantlanensis'' Grünwald, Reyes-Velasco, Franz-Chávez, Morales-Flores, Ahumada-Carrillo, Jones, and Boissinot, 2018 * '' E. (S.) marnockii'' Cope, 1878 * '' E. (S.) maurus'' Hedges, 1989 * '' E. (S.) modestus'' Taylor, 1942 * '' E. (S.) nietoi'' Grünwald, Reyes-Velasco, Franz-Chávez, Morales-Flores, Ahumada-Carrillo, Jones, and Boissinot, 2018 * '' E. (S.) nitidus'' Peters, 1870 * '' E. (S.) pallidus'' Duellman, 1958 * '' E. (S.) pipilans'' Taylor, 1940 * '' E. (S.) rubrimaculatus'' Taylor & Smith, 1945 * '' E. (S.) rufescens'' (Duellman & Dixon, 1959) * '' E. (S.) saxatilis'' Webb, 1962 * '' E. (S.) symingtoni'' Schwartz, 1957 * '' E. (S.) syristes'' Hoyt, 1965 * '' E. (S.) teretistes'' Duellman, 1958 * '' E. (S.) verrucipes'' Cope, 1885 * '' E. (S.) verruculatus'' Peters, 1870 * '' E. (S.) wixarika'' Reyes-Velasco, Ahumada-Carrillo, Burkhardt, and Devitt, 2015 * '' E. (S.) zeus'' Schwartz, 1958

References

{{Taxonbar, from=Q223286 Eleutherodactylinae Amphibian genera Taxa named by André Marie Constant Duméril Taxa named by Gabriel Bibron