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 14 F/ 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, 1 G ( 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 5 G ( 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 3 G ( 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) 4 G ( 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 1 G ( 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 3 G ( 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 1 G ( 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) 1 G ( 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) 3 G ( 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 1 G ( 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 1 G ( 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)