ANURA Hogg, 1839

Savage, Jay M. & Bolaños, Federico, 2009, A checklist of the Amphibians and Reptiles of Costa Rica: Additions and nomenclatural revisions, Zootaxa 2005, pp. 1-23 : 3-10

ORDER ANURA Hogg, 1839 14F/39+3*G (139) (136 + 3*)

The following family was included in the Dendrobatidae by Savage (2002). Grant et al. (2006) recognized it

as distinct from the latter family:

FAMILY AROMOBATIDAE Grant, Frost, Caldwell, Gagliardo, Haddad, Kok, Means, Noonan, Schargel , &

Wheeler, 2006, 1G (1)

Allobates Zimmermann & Zimmermann, 1988 (1)

Grant et al. (2006) recognized Allobates as distinct from Colostethu s. The following species was called Colostethu s talamancae by Savage (2002): Allobates talamancae (Cope, 1875) (381)

FAMILY BUFONIDAE J. E. Gray, 1825 5G (17) Atelopus C. Duméril & Bibron, 1841 (3)

Atelopus chiriquiensis Shreve, 1936 (187)

Atelopus senex Taylor, 1952 (188)

Atelopus varius ( Lichtenstein & von Martens, 1856) (189) Chaunus Wagler, 1828 (1)

Frost et al. (2006) recognized Chaunus (type species by monotypy: Chaunus marmoratus Wagler, 1828 = Bufo granulosus Spix, 1824 ) and Rhinella Fitzinger, 1826 (type species by monotypy: Oxyrhynchus proboscideus Spix, 1824 ) as genera distinct from Bufo . Chaparro, Pramuk and Gluesenkamp (2007) placed Chaunus as a synonym of Rhinella . Contrary to these authors implied statement that Chaunus is nested within Rhinella , this is not the case. Their phylogenetic tree shows that two monophyletic clades (sister groups) are subsumed within their concept of Rhinella . One of these includes the marinus , crucifer, and spinulosus groups and consitutes the genus Chaunus as used here. The other, Rhinella (sensu stricto), includes the margaritifer group of which Rhinella proboscidea is a member and a number of additional South American subclades.

The following species was called Bufo marinus in Savage (2002):

Chaunus marinus (Linné, 1758) (199)

Crepidophryne Cope, 1889 (3)

The following species were included in Crepidophryne epiotica by Savage (2002):

Crepidophryne chompipe Vaughan & Mendelson, 2007

Crepidophryne epiotica (Cope, 1875) (211)

Crepidophryne guanacaste Vaughan & Mendelson, 2007

Incilius Cope, 1863 (9)

Frost et al. (2006) recognized Cranopsis Cope, 1875 as distinct from Bufo . Inasmuch as Cranopsis is a junior homonym, Frost, Grant and Mendelson (2006) applied the name Ollotis to this genus. Unfortunately, the name Incilius Cope, 1863 (type species: Chilophryne conifera Cope, 1862 = Bufo coniferus Cope, 1862 by subsequent designation of Kellogg 1932) has priority over Ollotis (D. R. Frost, pers. comm.).

The following species were included in Bufo by Savage (2002):

The following species was included in Bufo melanochlorus by Savage (2002) but was recognized as

distinct by O’Neill & Mendelson (2004):

Incilius ancoinae ( O’Neill & Mendelson, 2004)

Incilius coccifer (Cope, 1866) (203)

Incilius coniferus (Cope, 1862) (207)

Incilius fastidiosus (Cope, 1875) (195)

Incilius holdridgei (Taylor, 1952) (196)

Incilius luetkenii (Boulenger, 1891) (208)

Incilius melanochlorus (Cope, 1877) (209)

Incilius periglenes (Savage, “1966”, 1967) (202)

Incilius valliceps (Wiegmann, 1833) (210)

Rhaebo Cope, 1862 (1)

Frost et al. (2006) recognized Rhaebo as distinct from Bufo .

The following species was called Bufo haematiticus by Savage (2002):

Rhaebo haematiticus Cope, 1862 (198)

FAMILY CENTROLENIDAE Taylor, 1951 3G (13) Centrolenella Noble, 1920 (2)

Centrolenella ilex Savage, 1967 (360)

Centrolenella prosoblepon (Boettger, 1892) (361) Cochranella Taylor, 1959 (5)

Cochranella albomaculata (Taylor, 1949) (365)

Cochranella euknemos (Savage & Starrett, 1967) (363)

Cochranella granulosa (Taylor, 1949) (363)

The following species was called Hyalinobatrachium pulveratum by Savage (2002) but was transferred to Cochranella by Cisneros-Heredia and McDiarmid (2006): Cochranella pulverata (W. Peters, 1873) (368)

Cochranella spinosa (Taylor, 1949) (365)

Hyalinobatrachium Ruiz and Lynch, 1991 (6)

Hyalinobatrachium chirripoi (Taylor, 1958) (369)

Hyalinobatrachium colymbiphyllum (Taylor, 1949) (370) Hyalinobatrachium fleischmanni (Boettger, 1893) (371)

Hyalinobatrachium talamancae (Taylor, 1952) (373)

Hyalinobatrachium valerioi (Dunn, 1931) (374)

Hyalinobatrachium vireovittatum (Starrett & Savage, 1973) (375)

Costa Rica members of the following family were referred to the family Leptodactylidae by Savage (2002) but Frost et al. (2006) placed them in the Brachycephalidae . Hedges et al. (2008) recognized them as distinct from both families.

FAMILY CRAUGASTORIDAE Hedges, Duellman & Heinicke, 2008 , 1G (28) Craugastor Cope, 1862 (28)

Crawford and Smith (2005) recognized Craugastor as distinct from Eleutherodactylus .

The following species were included in Eleutherodactylus by Savage (2002):

Craugastor andi (Savage, 1974) (237)

Craugastor angelicus (Savage, 1975) (248)

Craugastor bransfordii (Cope, “1885”, 1886) (257)

Craugastor catalinae (Campbell & Savage, 2000) (840)

Craugastor crassidigitus (Taylor, 1952) (238)

Craugastor cuaquero (Savage, 1980) (239)

Craugastor escoces (Savage, 1975) (249)

Craugastor fitzingeri ( O. Schmidt, 1857) (240)

Craugastor fleischmanni (Boettger, 1892) (250)

Craugastor gollmeri (W. Peters, 1863) (245)

Craugastor gulosus (Cope, 1875) (253)

Craugastor megacephalus (Cope, 1875) (254)

Craugastor melanostictus (Cope, 1875) (241)

Craugastor mimus (Taylor, 1955) (246)

Craugastor noblei (Barbour & Dunn, 1921) (247)

Craugastor obesus (Barbour, 1928) (840)

Craugastor persimilis (Barbour, 1926) (258)

Craugastor phasma (Lips & Savage, 1996) (242)

Craugastor podiciferus (Cope, 1875) (259)

Craugastor polyptychus (Cope, “1885”, 1886) (260)

Craugastor ranoides (Cope, “1885”, 1886) (251)

Craugastor rayo (Savage & DeWeese, 1979) (243)

Craugastor rhyacobatrachus (Campbell & Savage, 2000) (840)

Craugastor rugosus (W. Peters, 1873) (255)

Craugastor stejnegerianus (Cope, 1893) (261)

Craugastor talamancae (Dunn, 1931) (244)

Craugastor taurus (Taylor, 1958) (252)

Craugastor underwoodi (Boulenger, 1896) (262)

FAMILY DENDROBATIDAE Cope, 1865 (1850) 4G (7)

Dendrobates Wagler, 1830 (1)

Dendrobates auratus (Girard, “1854”, 1855) (383)

Oophaga Bauer, 1994 (2)

Grant et al. (2006) recognized Oophaga as distinct from Dendrobates . The following species were included in Dendrobates by Savage (2002): Oophaga granulifera (Taylor, 1958) (384)

Oophaga pumilio ( O. Schmidt, 1857) (386)

Phyllobates C. Duméril & Bibron, 1841 (2)

Phyllobates lugubris ( O. Schmidt, 1857) (388) Phyllobates vittatus (Cope, 1893) (390)

Silverstoneia Grant, Frost, Caldwell, Gagliardo, Haddad, Kok, Means, Noonan, Schargel , & Wheeler, 2006,

(2)

Grant et al. (2006) recognized Silverstoneia as distinct from Colostethu s. The following species were included in Colostethu s by Savage (2002): Silverstoneia flotator (Dunn, 1931) (378)

Silverstoneia nubicola (Dunn, 1924) (380)

Costa Rica members of the following family were referred to the family Leptodactylidae by Savage (2002) but Frost et al. (2006) placed them in the Brachycephalidae . Hedges et al. (2008) recognized them as distinct from both of these families.

FAMILY ELEUTHERODACTYLIDAE Lutz, 1954 1+1 G* (5) (4+1*) Diasporus Hedges, Duellman & Heinicke (2008) (4)

Heinicke et al. (2008) referred members of this genus to Pristimantis but Hedges et al. (2008)

recognized Diasporus as distinct from Eleutherodactylus and Pristimantis .

The following species were included in Eleutherodactylus by Savage (2002):

Diasporus diastema (Cope, 1875) (272)

Diasporus hylaeformis (Cope, 1875) (273)

Diasporus tigrillo (Savage, 1997) (274)

Diasporus vocator (Taylor, 1955) (274)

Eleutherodactylus C. Duméril & Bibron, 1841 (1 *)

The following species does not seem to be established in the country:

* Eleutherodactylus johnstonei Barbour, 1914 (271)

The family status of the following is unsettled. Frost et al. (2006) assigned the genus to the Amphignathodontidae Boulenger, 1882 . Wiens et al. (2007) placed it into a more inclusive Hemiphractidae Peters, 1862 . We have taken a conservation position and follow the latter authors. FAMILY HEMIPHRACTIDAE Peters, 1862 1G (1)

Gastrotheca Fitzinger, 1843 (1)

Gastrotheca cornuta (Boulenger, 1898) (294)

FAMILY HYLIDAE Rafinesque, 1815 15 + 1*G (42) (41+1 *) Subfamily HYLINAE Rafinesque, 1815 12 + 1*G (36) (35+1 *) Anotheca H. M.Smith, 1939 (1)

Anotheca spinosa (Steindachner, 1864) (295) Dendropsophus Fitzinger, 1843 (3)

Faivovich et al. (2005) recognized Dendropsophus as distinct from Hyla . The following species were included in Hyla by Savage (2002): Dendropsophus ebraccatus (Cope, 1874) (313)

Dendropsophus microcephalus (Cope, “1885”, 1886) (316) Dendropsophus phlebodes (Stejneger, 1906) (318)

Duellmanohyla Campbell & E. Smith, 1992 (3)

Duellmanohyla lythrodes (Savage, 1968) (297)

Duellmanohyla rufioculis (Taylor, 1952) (298)

Duellmanohyla uranochroa (Cope, 1875) (299)

Ecnomiohyla Faivovich, Haddad, Garcia, Frost, Campbell & Wheeler, 2005 , new genus (2)

Faivovich et al. (2005) recognized Ecnomiohyla as distinct from Hyla . The following species were included in Hyla by Savage (2002): Ecnomiohyla fimbrimembra (Taylor, 1948) (332) Ecnomiohyla miliaria (Cope, “1885”, 1886) (334)

Hyloscirtus W. Peters, 1882 (2)

Faivovich et al. (2005) recognized Hyloscirtus as distinct from Hyla . The following species were included in Hyla by Savage (2002): Hyloscirtus colymba (Dunn, 1931) (306)

Hyloscirtus palmeri (Boulenger, 1908) (307)

Hypsiboas Wagler, 1830 (2)

Faivovich et al. (2005) recognized Hypsiboas as distinct from Hyla . The following species were included in Hyla by Savage (2002): Hypsiboas rosenbergi (Boulenger, 1898) (303)

Hypsiboas rufitelus (Fouquette, 1961) (302)

Isthmohyla Faivovich, Haddad, Garcia, Frost, Campbell & Wheeler, 2005 (11)

Faivovich et al. (2005) recognized Isthmohyla as distinct from Hyla . The following species were included in Hyla by Savage (2002): Isthmohyla angustilineata (Taylor, 1952) (327)

Isthmohyla calypsa (Lips, 1996) (310)

Isthmohyla debilis (Taylor, 1952) (320)

Isthmohyla lancasteri (Barbour, 1928) (311)

Isthmohyla picadoi (Dunn, 1937) (336)

Isthmohyla pictipes (Cope, 1875) (321)

Isthmohyla pseudopuma (Günther, 1901) (328)

Isthmohyla rivularis (Taylor, 1952) (323)

Isthmohyla tica (Starrett, 1966) (324)

Isthmohyla xanthosticta (Duellman, 1968) (326) Isthmohyla zeteki (Gaige, 1929) (337)

* Osteopilus Fitzinger, 1843 (1)

* Osteopilus septentrionalis (C. Duméril & Bibron, 1841) (338)

Ptychohyla Taylor, 1944 (1)

Faivovich et al. (2005) recognized Ptychohyla as distinct from Hyla . The following species was called Hyla legleri by Savage (2002): Ptychohyla legleri (Taylor, 1958) (331)

Scinax Wagler, 1830 (3)

Scinax boulengeri (Cope, 1887) (343)

Scinax elaeochroa (Cope, 1875) (345)

Scinax staufferi (Cope, 1865) (346)

Smilisca Cope, 1865 (5)

Smilisca baudinii (C. Duméril & Bibron, 1841) (349) Smilisca phaeota (Cope, 1862) (351)

Smilisca puma (Cope, “1884”, 1885) (352)

Smilisca sila Duellman & Trueb, 1966 (354)

Smilisca sordida (W. Peters, 1863) (355)

Tlalocohyla Faivovich, Haddad, Garcia, Frost, Campbell & Wheeler, 2005 (1) Faivovich et al. (2005) recognized Tlalocohyla as distinct from Hyla . The following species was called Hyla loquax by Savage (2002): Tlalocohyla loquax (Gaige & Stuart, 1934) (308) Trachycephalus Tschudi, 1838 (1)

Faivovich et al. (2005) synonymized Phrynohyas with Trachycephalus . The following species was called Phrynohyas venulosus by Savage (2002): Trachycephalus venulosus (Laurenti, 1768) (340) Subfamily PHYLLOMEDUSINAE Günther, 1858 3G (6) Agalychnis Cope, 1864 (4)

Agalychnis annae (Duellman. 1963) (278)

Agalychnis callidryas (Cope, 1862) (281)

Agalychnis saltator Taylor, 1955 (283)

Agalychnis spurrelli Boulenger, 1913 (285)

Cruziohyla Faivovich, Haddad, Garcia, Frost, Campbell & Wheeler, 2005 (1) Faivovich et al. (2005) recognized Cruziohyla as distinct from Agalychnis . The following species was called Agalychnis calcarifer by Savage (2002): Cruziohyla calcarifer (Boulenger, 1902) (279)

Hylomantis W. Peters “1872”, 1873 (1)

Faviovich et al. (2005) recognized Hylomantis as distinct from Phyllomedusa . The following species was called Phyllomedusa lemur by Savage (2002): Hylomantis lemur (Boulenger, 1882) (286)

The members of the following family were included in the Leptodactylidae by Savage (2002). Frost et al.

(2006) recognized it as distinct from the latter family.

FAMILY LEIUPERIDAE Bonaparte, 1850 1G (1)

Engystomops Jiménez de la Espada, 1872 (1)

Nascimento, Caramaschi, and Cruz (2005), recognized Engystomops as distinct from Physalaemus . The following species was called Physalaemus pustulosus by Savage (2002): Engystomops pustulosus (Cope, 1864) (224)

FAMILY LEPTODACTYLIDAE Werner, 1896 (1838) 1G (5)

Leptodactylus Fitzinger, 1826 (5)

The following species was called Leptodactylus labialis by Savage (2002). Heyer (2002) recognized it as a distinct species:

Leptodactylus fragilis (Brocchi, 1877) (221)

The following species was called Leptodactylus bolivianus by Savage (2002) but is recognized as a distinct species by W. R. Heyer (pers. comm.):

Leptodactylus insularum Barbour, 1906 (217)

Leptodactylus melanonotus (Hallowell, “1860”, 1861) (215) Leptodactylus poecilochilus (Cope, 1862) (222)

The following species was included in Leptodactylus pentadactylus by Savage (2002): Leptodactylus savagei Heyer, 2005 (219)

FAMILY MICROHYLIDAE Günther, 1858 (1843) 3G (3) Gastrophryne Fitzinger, 1843 (1)

Gastrophryne pictiventris (Cope, “1885”, 1886) (393)

Hypopachus Keferstein, 1867 (1)

Hypopachus variolosus (Cope, 1866) (394) Nelsonophryne Frost, 1987 (1)

Nelsonophryne aterrima (Günther, 1900) (396)

FAMILY RANIDAE Rafinseque, 1814 1G (6) (5+1*)

Lithobates Fitzinger, 1843 (6) (5+1 *)

Frost et al. (2006) recognized Lithobates as distinct from Rana a result confirmed by Che et al. (2007). The following species does not seem to be established in the country: * Lithobates catesbeianus (Shaw, 1802) , addition to fauna The following species were included in Rana by Savage (2002): Lithobates forreri (Boulenger, 1883) (399)

Lithobates taylori (H. M. Smith, 1959) (400)

Lithobates vaillanti (Brocchi, 1877) (402)

Lithobates vibicarius (Cope, 1894) (403)

Lithobates warszewitschii ( O. Schmidt, 1857) (404)

FAMILY RHINOPHRYNIDAE Günther , “1858”, 1859 1G (1) Rhinophrynus C. Duméril & Bibron, 1841 (1)

Rhinophrynus dorsalis C. Duméril & Bibron, 1841 (183)

Costa Rica members of the following family were referred to the family Leptodactylidae by Savage (2002) but Frost et al. (2006) placed them in the Brachycephalidae . Hedges et al. (2008) recognized them as distinct from both of these families.

STRABOMANTIDAE Hedges, Duellman & Henicke, 2008 , 2G (9) Pristimantis Jiménez de la Espada, “1870”, 1871 (8)

Heinicke, Duellman, and Hedges (2007) recognized Pristimantis as distinct from both Craugator and Eleutherodactylus .

The following species were included in Eleutherodactylus by Savage (2002):

Pristimantis altae (Dunn, 1942) (265)

Pristimantis caryophyllaceus (Barbour, 1928) (266)

Pristimantis cerasinus (Cope, 1875) (264)

Pristimantis cruentus (W. Peters, 1873) (267)

Pristimantis gaigeae (Dunn, 1931) (263)

Pristimantis moro (Savage, 1965) (268)

Pristimantis pardalis (Barbour, 1928) (269)

Pristimantis ridens (Cope, 1866) (269)

Strabomantis Peters, 1863 (1)

Heinicke et al. (2006) recognized Lymnophis Jiménez de la Espada, “1870”, 1871 as distinct from Craugastor and Eleutherodactylus . Inasmuch as Limnophys is a junior homonym, Hedges et al. (2008) applied the name Strabomantis to this genus.

The following species was included in Eleutherodactylus by Savage (2002): Strabomantis bufoniformis (Boulenger, 1896) (256)