Cliff chirping frog, Eleutherodactylus marnockii
Eleutherodactylus is a genus of frogs in the family Eleutherodactylidae. Many of the 185 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 (E. coqui), which is both a national symbol of Puerto Rico and a notorious invasive species 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 eleutheros (ἐλεύθερος, ‘free, unbound’) and dactylos (δάκτυλος, ‘finger, toe’).

Most species are small, slender, and cryptically colored, with three to five free toes. A few, such as the web-footed coquí (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 (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.

Phylogenetics

The basis of forming this genus has been morphological, but sequence comparisons of protein-encoding DNA, mitochondrial DNA, and ribosomal RNA 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. All true members of the genus have been clustered into subgenera, 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 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 epochs. Land bridges would have been limited to facilitating dispersal between West Indian islands, however, the Oligocene division of Hispaniola 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 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, however, it should exclude clades with distributions south of the Panama Canal.

Species

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
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.) 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.) winlandi Barbour, 1914
E. (Eu.) zugi Schwartz, 1958
E. (P.) chlorophenax Schwartz, 1976
E. (P.) hypostenor Schwartz, 1965
E. (P.) inoptatus Barbour, 1914
E. (P.) nortoni Schwartz, 1976
E. (P.) parapelates Hedges & Thomas, 1987
E. (P.) ruthae Noble, 1923
E. (S.) albolabris Lynch & Lescure, 1980
E. (S.) angustidigitorum Taylor, 1940
E. (S.) cystingnathoides Cope, 1877
E. (S.) dennisi Lynch, 1970
E. (S.) dilatus Davis & Dixon, 1955
E. (S.) grandis Dixon, 1957
E. (S.) guttilatus Cope, 1879
E. (S.) leprus Cope, 1879
E. (S.) interorbitalis Langebartel & Shannon, 1956
E. (S.) longipes Baird, 1859
E. (S.) marnockii Cope, 1878
E. (S.) maurus Hedges, 1989
E. (S.) modestus Taylor, 1942
E. (S.) nitidus Peters, 1870
E. (S.) nivicolimae Dixon & Webb, 1966
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.) zeus Schwartz, 1958
E. adamastus Campbell, 1994
E. alfredi Boulenger, 1898
E. amniscola Campbell & Savage, 2000
E. anciano Savage, McCranie & Wilson, 1988
E. andi Savage, 1974
E. angelicus Savage, 1975
E. aphanus Campbell, 1994
E. augusti Dugès, 1879
E. aurilegulus Savage, McCranie & Wilson, 1988
E. azueroensis Savage, 1975
E. batrachylus Taylor, 1940
E. berkenbuschii Peters, 1870
E. biporcatus Peters, 1863
E. bocourti Brocchi, 1877
E. bransfordii Cope, 1886
E. brocchi Boulenger, 1882
E. campbelli Smith, 2005
E. catalinae Campbell & Savage, 2000
E. chac Savage, 1987
E. charadra Campbell & Savage, 2000
E. chrysozetetes McCranie, Savage & Wilson, 1989
E. coffeus McCranie & Köhler, 1999
E. crassidigitus Taylor, 1952
E. cruzi McCranie, Savage & Wilson, 1989
E. cuaquero Savage, 1980
E. cyanocthebius McCranie & Smith, 2006
E. daryi Ford & Savage, 1984
E. latens Lynch, 1989
E. decoratus Taylor, 1942
E. emcelae Lynch, 1985
E. emleni Dunn & Emlen, 1932
E. epochthidius McCranie & Wilson, 1997
E. escoces Savage, 1975
E. fecundus McCranie & Wilson, 1997
E. fitzingeri Schmidt, 1857
E. fleischmanni Boettger, 1892
E. galacticorhinus Canseco-Marquez & Smith, 2004
E. glaucus Lynch, 1967
E. gollmeri Peters, 1863
E. greggi Bumzahem, 1955
E. guerreroensis Lynch, 1967
E. gulosus Cope, 1875
E. hobartsmithi Taylor, 1937
E. inachus Campbell & Savage, 2000
E. jota Lynch, 1980
E. laevissimus Werner, 1896
E. laticeps Duméril, 1853
E. lauraster Savage, McCranie & Espinal, 1996
E. lineatus Brocchi, 1879
E. loki Shannon & Werler, 1955
E. longirostris Boulenger, 1898
E. matudai Taylor, 1941
E. megacephalus Cope, 1875
E. megalotympanum Shannon & Werler, 1955
E. melanostictus Cope, 1875
E. merendonensis Schmidt, 1933
E. mexicanus Brocchi, 1877
E. milesi Schmidt, 1933
E. mimus Taylor, 1955
E. monnichorum Dunn, 1940
E. montanus Dunn, 1940
E. myllomyllon Savage, 2000
E. nefrens Smith, 2005
E. noblei Barbour & Dunn, 1921
E. obesus Barbour, 1928
E. occidentalis Taylor, 1941
E. olanchano McCranie & Wilson, 1999
E. omiltemanus Günther, 1900
E. omoaensis McCranie & Wilson, 1997
E. opimus Savage & Myers, 2002
E. palenque Campbell & Savage, 2000
E. pechorum McCranie & Wilson, 1999
E. pelorus Campbell & Savage, 2000
E. persimilis Barbour, 1926
E. phasma Lips & Savage, 1996
E. podiciferus Cope, 1875
E. polymniae Campbell, Lamar & Hillis, 1989
E. polyptychus Cope, 1886
E. pozo Johnson & Savage, 1995
E. psephosypharus Campbell, Savage & Meyer, 1994
E. punctariolus Peters, 1863
E. pygmaeus Taylor, 1937
E. raniformis Boulenger, 1896
E. ranoides Cope, 1886
E. rayo Savage & DeWeese, 1979
E. rhodopsis Cope, 1867
E. rhyacobatrachus Campbell & Savage, 2000
E. rivulus Campbell & Savage, 2000
E. rostralis Werner, 1896
E. rugosus Peters, 1873
E. rugulosus Cope, 1870
E. rupinius Campbell & Savage, 2000
E. sabrinus Campbell & Savage, 2000
E. saltuarius McCranie & Wilson, 1997
E. sandersoni Schmidt, 1941
E. silvicola Lynch, 1967
E. spatulatus Smith, 1939
E. stadelmani Schmidt, 1936
E. stejnegerianus Cope, 1893
E. stuarti Lynch, 1967
E. tabasarae Savage, Hollingsworth, Lips & Jaslow, 2004
E. talamancae Dunn, 1931
E. tarahumaraensis Taylor, 1940
E. taurus Taylor, 1958
E. taylori Lynch, 1966
E. trachydermus Campbell, 1994
E. underwoodi Boulenger, 1896
E. uno Savage, 1984
E. vocalis Taylor, 1940
E. vulcani Shannon & Werler, 1955
E. xucanebi Stuart, 1941
E. yucatensis Lynch, 1965

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