Bufonidae

North Borneo stream toad Ansonia fuliginea, copyright Ivan Neo.

Belongs within: Hyloidea.
Contains: Atelopus, Rhinella, Cranopsis, Bufo, Amietophrynus.

The Bufonidae, true toads, are a cosmopolitan family of frogs with a typically warty appearance.

Synapomorphies (from Frost et al. 2006): Larva with diastema in larval lower lip papillation; m. diaphragmatopraecordialis absent; lateral fibers of m. subarcualis rectus II–IV invading interbranchial septum IV; processus anterolateralis of crista parotica absent; larval lungs rudimentary or absent.

<==Bufonidae [Bufonina]
|--Melanophryniscus Gallardo 1961PW11, FG06
| | i. s.: M. moreiraeCER68
| | M. orejasmirandaiFG06
| |--M. rubriventrisPW11
| `--+--M. klappenbachiPW11
| `--M. stelzneriPW11
| |--M. s. stelzneriCER68
| `--M. s. ateoluteusCER68
`--+--AtelopusPW11
`--+--Osornophryne Ruiz-Carranza & Hernández-Camacho 1976PW11, FG06
| |--O. sumacoensisPW11
| `--+--O. guacamayoPW11
| `--+--O. puruantaPW11
| `--+--O. antisanaPW11
| `--O. bufoniformisPW11
`--+--+--Dendrophryniscus Jiménez de la Espada 1871PW11, FG06 [Dendrophryniscina]
| | |--D. brevipillicatusNW07
| | `--D. minutusPW11
| `--Oreophrynella Boulenger 1895NW07, FG06
| |--O. crypticaVSD02
| |--O. huberiVSD02
| |--O. quelchiiNW07
| `--O. vasqueziVSD02
`--+--+--‘Bufo’ cophotis Boulenger 1900PW11, FG06 [=Chaunus cophotisFG06]
| `--Nannophryne Günther 1870FG06
| |--*N. variegata Günther 1870FG06 [=Bufo variegatusPW11]
| `--N. cophotisFB17
`--+--+--+--‘Bufo’ nasicus Werner 1903PW11, FG06 [=Rhinella nasicaFG06]
| | `--Rhaebo Cope 1862FB17, FG06 [incl. Phrynomorphus Fitzinger 1843 non Curtis 1831FG06]
| | | i. s.: R. nasicusFB17
| | |--*R. haematiticus (Cope 1862)FG06 [=Bufo haematiticusPW11]
| | `--+--B. glaberrimus (Günther 1869)FG06 [=Bufo glaberrimusPW11]
| | `--R. guttatus (Schneider 1799)FG06 (see below for synonymy)
| `--Peltophryne Fitzinger 1843FB17, FG06
| |--P. lemur (Cope 1869)FG06 [=Bufo lemurPW11]
| `--+--P. guentheri (Cochran 1941)FG06 [=Bufo guentheriPW11]
| `--+--+--P. gundlachi (Ruibal 1959)FG06 [=Bufo gundlachiPW11]
| | `--P. longinasus (Stejneger 1905)FG06 [=Bufo longinasusPW11]
| `--+--P. taladai (Schwartz 1960)FG06 [=Bufo taladaiPW11]
| `--+--P. empusa Cope 1862FG06 [=Bufo empususPW11]
| `--+--P. fustiger (Schwartz 1960)FG06 [=Bufo peltocephalus fustigerPHH01, PW11]
| `--*P. peltocephala (Tschudi 1838)FG06 [=Bufo peltocephalusPW11]
`--+--+--RhinellaPW11
| `--+--‘Cranopsis’PW11
| `--+--Anaxyrus Tschudi 1845FB17, FG06
| | | i. s.: A. compactilis (Wiegmann 1833) (see below for synonymy)FG06
| | |--A. punctatus (Baird & Girard 1852)FG06 [=Bufo punctatusPW11]
| | `--+--+--A. boreas (Baird & Girard 1852)FG06 [=Bufo boreasPW11]
| | | | |--A. b. boreasCER68
| | | | `--‘Bufo’ b. halophilusCER68
| | | `--A. nelsoni (Stejneger 1893)FG06 [=Bufo nelsoniPW11]
| | `--+--A. canorus (Camp 1916)FB17, FG06 [=Bufo canorusFG06]
| | `--A. exsul (Myers 1942)FG06 [=Bufo exsulPW11]
| `--+--‘Bufo’ quercicus Holbrook 1840PW11, FG06 [=Anaxyrus quercicusFG06]
| `--+--+--+--‘Bufo’ debilis Girard 1854PW11, FG06 [=Anaxyrus debilisFG06]
| | | `--‘Bufo’ retiformis Sanders & Smith 1951PW11, FG06 [=Anaxyrus retiformisFG06]
| | `--Incilius Cope 1863FB17, FG06
| | |--*I. cognatus (Say in James 1823)FG06 [=Bufo cognatusPW11, Anaxyrus cognatusFG06]
| | |--I. luetkeniiFB17
| | `--I. nebuliferFB17
| `--+--‘Bufo’ speciosus Girard 1854PW11, FG06 (see below for synonymy)
| `--+--‘Bufo’ californicus Camp 1915PW11, FG06 [=Anaxyrus californicusFG06]
| `--+--‘Bufo’ microscaphus Cope 1867PW11, FG06 [=Anaxyrus microscaphusFG06]
| `--+--‘Bufo’ fowleri Hinckley 1882PW11, FG06 (see below for synonymy)
| `--+--‘Bufo’ terrestris Bonnaterre 1789PW11, FG06 [=Anaxyrus terrestrisFG06]
| `--+--‘Bufo’ woodhousii Girard 1854PW11, FG06 (see below for synonymy)
| | |--B. w. woodhousiiCER68
| | `--B. w. australisCER68
| `--+--‘Bufo’ baxteri Porter 1968PW11, FG06 [=Anaxyrus baxteriFG06]
| `--+--‘Bufo’ hemiophrys Cope 1886PW11, FG06 [=Anaxyrus hemiophrysFG06]
| `--+--‘Bufo’ americanus Holbrook 1836PW11, FG06 (see below for synonymy)
| | |--B. a. americanusPHC04
| | `--B. a. charlesmithiPHC04
| `--‘Bufo’ houstonensis Sanders 1953PW11, FG06 (see below for synonymy)
`--+--+--+--Schismaderma Smith 1849FB17, FG06
| | | `--S. carensPW11
| | `--+--+--‘Bufo’ boulengeriPW11
| | | `--‘Bufo’ siculusPW11
| | `--Pseudepidalea Frost, Grant et al. 2006FG06
| | |--+--P. oblonga (Nikolskii 1896)FG06 [=Bufo oblongusPW11]
| | | | |--P. o. oblongaFG06
| | | | `--P. o. danatensis [=Bufo oblongus danatensis]FG06
| | | `--P. pewzowi (Bedriaga 1898)FG06 [=Bufo pewzowiPW11]
| | `--+--‘Bufo’ balearicusPW11
| | `--+--*P. viridis (Laurenti 1768)FG06 [=Bufo viridisPW11, Rana viridisK08]
| | `--‘Bufo’ variabilisPW11
| `--+--+--Churamiti Channing & Stanley 2002PW11, FG06
| | | `--C. maridadiPW11
| | `--Nectophrynoides Noble 1926FB17, FG06
| | | i. s.: N. cryptusFG06
| | |--N. viviparusPW11
| | `--+--N. minutusPW11
| | `--N. tornieriPW11
| `--+--Phrynoidis Fitzinger 1843FB17, FG06
| | |--*P. aspera (Gravenhorst 1829)FG06 [=Bufo asperPW11]
| | `--P. juxtaspera (Inger 1964)FG06 [=Bufo juxtasperPW11]
| `--Pedostibes Günther 1876PW11, FG06
| |--P. hosiiPW11
| |--P. rugosusPW11
| `--P. tuberculosusPW11
`--+--Rentapia hosiiFB17
`--+--+--‘Bufo’ brongersmai Hoogmoed 1972PW11, FG06 [=Pseudepidalea brongersmaiFG06]
| `--+--Adenomus Cope 1861PW11, FG06 [Adenomidae]
| | `--A. kelaartiiPW11
| `--+--+--‘Bufo’ koynayensis Soman 1963PW11, FG06
| | `--+--‘Bufo’ dhufarensis Parker 1931PW11, FG06
| | `--+--‘Bufo’ hololius Günther 1876PW11, FG06
| | `--‘Bufo’ stomaticus Lütken 1864PW11, FG06
| `--Duttaphrynus Frost, Grant et al. 2006FG06
| | i. s.: D. stomaticusFB17
| |--D. crocus (Wogan, Win et al. 2003)FG06 [=Bufo crocusPW11]
| `--+--‘Bufo’ stuarti Smith 1929PW11, FG06
| `--+--D. himalayanus (Günther 1864)FG06 [=Bufo himalayanusPW11]
| `--+--+--‘Bufo’ atukoralei Bogert & Senanayake 1966PW11, FG06
| | `--‘Bufo’ scaber Schneider 1799PW11, FG06
| `--+--*D. melanostictus (Schneider 1799)FG06 [=Bufo melanostictusPW11]
| `--+--‘Bufo’ brevirostris Rao 1937PW11, FG06
| `--‘Bufo’ parietalis Boulenger 1882PW11, FG06
`--+--BufoFB17
|--+--Epidalea Cope 1864FB17, FG06
| | `--*E. calamita (Laurenti 1768)FG06 [=Bufo calamitaPW11]
| `--Leptophryne Fitzinger 1843PW11, FG06
| `--L. borbonicaPW11
`--+--+--+--Sabahphrynus maculatusPW11
| | `--Ingerophrynus Frost, Grant et al. 2006FB17, FG06
| | |--I. celebensis (Günther 1859)FG06 [=Bufo celebensisPW11]
| | |--+--*I. biporcatus (Gravenhorst 1829)FG06 [=Bufo biporcatusPW11]
| | | `--I. divergens (Peters 1871)FG06 [=Bufo divergensPW11]
| | `--+--I. galeatus (Günther 1864)FG06 [=Bufo galeatusPW11]
| | |--I. macrotis (Boulenger 1887)FG06 [=Bufo macrotisPW11]
| | `--I. philippinicus (Boulenger 1887)FG06 [=Bufo philippinicusPW11]
| `--+--+--‘Ansonia’ ornataPW11
| | `--Didynamipus Andersson 1903PW11, FG06
| | `--D. sjostedtiPW11
| `--+--Pelophryne Barbour 1938PW11, FG06
| | |--P. miseraPW11
| | `--+--P. brevipesPW11
| | `--P. signataPW11
| `--Ansonia Stoliczka 1870PW11, FG06
| |--+--+--A. fuligineaPW11
| | | `--A. muelleriPW11
| | `--+--A. leptopusPW11
| | `--A. longidigitaPW11
| `--+--A. malayanaPW11
| `--+--A. spinuliferPW11
| `--+--A. hanitschiPW11
| `--+--A. minutaPW11
| `--A. platysomaPW11
`--+--+--Capensibufo Grandison 1980PW11, FG06
| | |--C. roseiPW11
| | `--C. tradouwiPW11 [=Bufo tradouwiFG06]
| `--+--‘Bufo’ mauritanicus Schlegel 1841PW11, FG06
| `--AmietophrynusPW11
`--+--Vandijkophrynus Frost, Grant et al. 2006FG06
| |--*V. angusticeps (Smith 1848)FG06 [=Bufo angusticepsPW11]
| `--+--+--V. amatolicus (Hewitt 1925)FG06 [=Bufo amatolicusPW11]
| | `--V. inyangae (Poynton 1963)FG06 [=Bufo inyangaePW11]
| `--+--V. gariepensis (Smith 1848)FG06 [=Bufo gariepensisPW11]
| `--V. robinsoni (Branch & Braacke 1996)FG06 [=Bufo robinsoniPW11]
`--+--+--‘Bufo’ fenoulheti Hewitt & Methuen 1912PW11, FG06 (see below for synonymy)
| | |--B. f. fenoulhetiM54
| | |--B. f. obtususM54
| | `--B. f. rhodesianusM54
| `--+--‘Bufo’ damaranus Mertens 1954PW11, FG06 (see below for synonymy)
| `--‘Bufo’ dombensis Bocage 1895PW11, FG06 [=Poyntonophrynus dombensisFG06]
`--+--+--‘Bufo’ taitanus Peters 1878PW11, FG06 [=Mertensophryne taitanaFG06]
| `--‘Bufo’ uzunguensis Loveridge 1932PW11, FG06 (see below for synonymy)
`--+--+--‘Bufo’ lindneri Mertens 1955PW11, FG06 [=Mertensophryne lindneriFG06]
| `--Stephopaedes Channing 1979PW11, FG06 [Stephopaedini]
| |--S. anotis (Boulenger 1907) (see below for synonymy)FG06
| |--S. howelli Poynton & Clarke 1999 (see below for synonymy)FG06
| |--S. loveridgei (Poynton 1991) (see below for synonymy)FG06
| `--S. usambarae (Poynton & Clarke 1999) (see below for synonymy)FG06
`--Mertensophryne Tihen 1960FB17, FG06
|--*M. rondoensis (Loveridge 1942) [=Bufo micranotis rondoensis]FG06
|--M. lonnbergi (Andersson 1911) [=Bufo lonnbergi]FG06
|--M. melanopleura (Schmidt & Inger 1959) [=Bufo melanopleurus]FG06
|--M. micranotis (Loveridge 1925) [=Bufo micranotis]FG06
|--M. mocquardi (Angel 1924) [=Bufo mocquardi]FG06
|--M. nyikae (Loveridge 1953) [=Bufo nyikae]FG06
`--M. schmidti (Grandison 1972) [=Bufo schmidti]FG06

Bufonidae incertae sedis:
Altiphrynoides Dubois 1987 [incl. Spinophrynoides Dubois 1987]FG06
|--A. malcolmiFG06
`--A. osgoodi (Loveridge 1932) [=Spinophrynoides osgoodi]FG06
Truebella Graybeal & Cannatella 1995FG06
Pseudobufo Tschudi 1838FG06
Crepidophryne Cope 1889FG06
`--C. epioticaFG06
Andinophryne Hoogmoed 1985FG06
Atelophryniscus McCranie, Wilson & Williams 1989FG06
Bufoides Pillai & Yazdani 1983FG06
Frostius Cannatella 1986FG06
Laurentophryne Tihen 1960FG06
`--L. parkeriFG06
Metaphryniscus Señaris, Ayarzagüena & Gorzula 1994FG06
Nimbaphrynoides Dubois 1987FG06
|--N. liberiensisFG06
`--N. occidentalisFG06
Parapelophryne Fei, Ye & Jiang 2003FG06
Phryniscus Wiegmann 1834 [Phryniscidae]FG06
`--P. nigricansB89
Platosphus d’Isle 1877 [Platosphinae]FG06
Bufavus Portis 1885 [Bufavidae]FG06
`--B. meneghinii Ports 1885RR00
Tornierobates Miranda-Ribeiro 1926 [Tornierobatidae]FG06
Amazophrynella minutaFB17
SclerophrysFB17
|--S. brauniFB17
`--S. maculataFB17
Bufotes viridisFB17
0--Nectophryne Buchholz & Peters 1875PW11, FG06 [Nectophrynidae]
| |--N. afraPW11
| `--N. batesiiPW11
`--+--Werneria Poche 1903PW11, FG06
| |--W. mertensianaPW11
| `--W. preussiFG06
`--Wolterstorffina Mertens 1939PW11, FG06
|--W. mireiFG06
`--W. parvipalmataPW11

Anaxyrus compactilis (Wiegmann 1833) [=Bufo compactilis; incl. *A. melancholicus Tschudi 1845]FG06

‘Bufo’ americanus Holbrook 1836PW11, FG06 [=Anaxyrus americanusFG06, B. terrestris americanusCM07]

‘Bufo’ damaranus Mertens 1954PW11, FG06 [=B. fenoulheti damaranusM54, Poyntonophrynus damaranusFG06]

‘Bufo’ fenoulheti Hewitt & Methuen 1912PW11, FG06 [=Poyntonophrynus fenoulhetiFG06]

‘Bufo’ fowleri Hinckley 1882PW11, FG06 [=Anaxyrus fowleriFG06, B. woodhousei fowleriCM07]

‘Bufo’ houstonensis Sanders 1953PW11, FG06 [=Anaxyrus houstonensisFG06]

‘Bufo’ speciosus Girard 1854PW11, FG06 [=Anaxyrus speciosusFG06, B. compactilis speciosusCM07]

Bufo’ uzunguensis Loveridge 1932PW11, FG06 [=Mertensophryne uzunguensisFG06]

‘Bufo’ woodhousii Girard 1854PW11, FG06 [=Anaxyrus woodhousiiFG06, B. lentiginosus woodhouseiDS89; incl. B. woodhousii bexarensis Mecham 1959H00]

Rhaebo guttatus (Schneider 1799)FG06 [=Bufo guttatusPW11; incl. B. leschenaulti Duméril & Bibron 1841FG06, *Phrynomorphus leschenaultiFG06]

Stephopaedes anotis (Boulenger 1907) [=Mertensophryne (Stephopaedes) anotis]FG06

Stephopaedes howelli Poynton & Clarke 1999 [=Mertensophryne (Stephopaedes) howelli]FG06

Stephopaedes loveridgei (Poynton 1991) [=Mertensophryne (Stephopaedes) loveridgei]FG06

Stephopaedes usambarae (Poynton & Clarke 1999) [=Mertensophryne (Stephopaedes) usambarae]FG06

*Type species of generic name indicated

References

[B89] Boulenger, G. A. 1889. On a collection of batrachians made by Prof. Charles Spegazzini at Colonia Resistencia, South Chaco, Argentine Republic. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 246–249.

[CER68] Cei, J. M., V. Erspamer & M. Roseghini. 1968. Taxonomic and evolutionary significance of biogenic amines and polypeptides in amphibian skin. II. Toads of the genera Bufo and Melanophryniscus. Systematic Zoology 17 (3): 232–245.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[DS89] Dundee, H. A., & H. M. Smith. 1989. Nomenclatural eponym endings revisited. Systematic Zoology 38 (3): 279–283.

[FB17] Feng, Y.-J., D. C. Blackburn, D. Liang, D. M. Hillis, D. B. Wake, D. C. Cannatella & P. Zhang. 2017. Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary. Proceedings of the National Academy of Sciences of the USA 114 (29): E5864–E5870.

[FG06] Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. Haas, C. F. B. Haddad, R. O. de Sá, A. Channing, M. Wilkinson, S. C. Donnellan, C. J. Raxworthy, J. A. Campbell, B. L. Blotto, P. Moler, R. C. Drewes, R. A. Nussbaum, J. D. Lynch, D. M. Green & W. C. Wheeler. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1–370.

[H00] Holman, J. A. 2000. Pleistocene Amphibia: evolutionary stasis, range adjustments, and recolonization patterns. In: Heatwole, H., & R. L. Carroll (eds) Amphibian Biology vol. 4. Palaeontology. The evolutionary history of amphibians pp. 1445–1458. Surrey Beatty & Sons.

[K08] Korsós, Z. 2008. History of the Herpetological Collection of the Hungarian Natural History Museum. Annales Historico-Naturales Musei Nationalis Hungarici 100: 37–93.

[M54] Mertens, R. 1954. Eine neue Kröte aus Südwestafrika. Senckenbergiana Biologica 35 (1–2): 9–11.

[NW07] Nussbaum, R. A., & S.-H. Wu. 2007. Morphological assessments and phylogenetic relationships of the Seychellean frogs of the family Sooglossidae (Amphibia: Anura). Zoological Studies 46 (3): 322–335.

[PHC04] Pauly, G. B., D. M. Hillis & D. C. Cannatella. 2004. The history of a Nearctic colonization: Molecular phylogenetics and biogeography of the Nearctic toads (Bufo). Evolution 58 (11): 2517–2535.

[PHH01] Pramuk, J. B., C. A. Hass & S. B. Hedges. 2001. Molecular phylogeny and biogeography of West Indian toads (Anura: Bufonidae). Molecular Phylogenetics and Evolution 20 (2): 294–301.

[PW11] Pyron, R. A., & J. J. Wiens. 2011. A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution 61: 543–583.

[RR00] Roček, Z., & J.-C. Rage 2000. Tertiary Anura of Europe, Africa, Asia, North America, and Australia. In: Heatwole, H., & R. L. Carroll (eds) Amphibian Biology vol. 4. Palaeontology. The evolutionary history of amphibians pp. 1332–1387. Surrey Beatty & Sons.

[VSD02] Villarreal, O., C. Señaris & C. DoNascimiento. 2002. Contribución al conocimiento faunístico del Wei-Assipu-tepui, macizo del Roraima, con énfasis en la anurofauna y opiliofauna. Boletín de la Sociedad Venezolana de Espeleología 36: 46–50.

Leave a comment

Your email address will not be published. Required fields are marked *