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Lithobates; Temporal range: Early Miocene–present PreꞒ Ꞓ O S D C P T J K Pg N
	A close up of a male American bullfrog (Lithobates catesbeianus)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Amphibia
Order:	Anura
Family:	Ranidae
Genus:	Lithobates; Fitzinger, 1843
Type species
	Rana palmipes; Spix, 1824
Species
	7 to 50, depending on the definition
Synonyms
	†Anchylorana Taylor, 1942;

Lithobates, commonly known as the bullfrogs, is a genus of true frogs, of the family Ranidae.^[1] The name is derived from litho- (stone) and the Greek bates (βάτης, one that treads), meaning one that treads on rock, or rock climber.^[2]^[3] As presently defined, it includes many of eastern North America's most familiar aquatic frog species, including the American bullfrog, green frog, and the leopard frogs.

Systematics

The name was defined by Hillis and Wilcox (2005) for a subgenus of four Central and South American frogs within the genus Rana.^[4]^[5] The subgenus was subsequently expanded to seven species in Central and South America in a systematic revision of the genus Rana.^[6] The name was previously used by Frost et al. as a separate genus of ranid frogs that included most of the North American frogs traditionally included in the genus Rana,^[7] including the American bullfrog and northern leopard frog. Frost used the name in this sense in the frog section of a North American common names list edited by Crother (2008).^[8] This proposed change has since been rejected by others, such as Stuart (2008),^[9] Pauly et al. (2009),^[10] AmphibiaWeb,^[11] and Yuan et al. (2016).^[6] AmphibiaWeb, available at http://amphibiaweb.org/, an online compendium of amphibian names, follows Yuan et al. (2016) in recognizing Lithobates as a subgenus.^[6] On the other hand, Amphibian Species of the World 6.0, an online reference, uses Lithobates as a genus.^[1] This definition is also followed by, e.g., the International Union for Conservation of Nature (IUCN)^[12] and the Society for the Study of Amphibians and Reptiles.^[13]

The earliest known members of this genus are known from the Early Miocene of Florida, and appear to belong to the leopard frog species complex.^[14]

Species

Recent species

These species are recognised in the genus Lithobates:^[1]

Lithobates areolatus (Baird and Girard, 1852) – crawfish frog
Lithobates berlandieri (Baird, 1859) – Rio Grande leopard frog
Lithobates blairi (Mecham, Littlejohn, Oldham, Brown, and Brown, 1973) – Plains leopard frog
Lithobates brownorum (Sanders, 1973)
Lithobates bwana (Hillis and de Sá, 1988) – Rio Chipillico frog
Lithobates capito (LeConte, 1855) – gopher frog
Lithobates catesbeianus (Shaw, 1802) – American bullfrog
Lithobates chichicuahutla (Cuellar, Méndez-De La Cruz, and Villagrán-Santa Cruz, 1996) – Lago de las Minas frog
Lithobates chiricahuensis (Platz and Mecham, 1979) – Chiricahua leopard frog
Lithobates clamitans (Latreille, 1801) – green frog
Lithobates dunni (Zweifel, 1957) – Patzcuaro frog
†Lithobates fisheri (Stejneger, 1893) – Vegas Valley leopard frog or Mogollon Rim leopard frog
Lithobates forreri (Boulenger, 1883) – Forrer's leopard frog
Lithobates grylio (Stejneger, 1901) – pig frog
Lithobates heckscheri (Wright, 1924) – river frog
Lithobates johni (Blair, 1965) – Moore's frog
Lithobates juliani (Hillis and de Sá, 1988) – Maya Mountains frog
Lithobates kauffeldi (Feinberg, Newman, Watkins-Colwell, Schlesinger, Zarate, Curry, Shaffer, and Burger, 2014) – Atlantic Coast leopard frog
Lithobates lenca (Luque-Montes et al., 2018)
Lithobates lemosespinali (Smith and Chiszar, 2003) – Lemos-Espinal's leopard frog
Lithobates macroglossa (Brocchi, 1877) – Guatemala plateau frog
Lithobates maculatus (Brocchi, 1877) – highland frog
Lithobates magnaocularis (Frost and Bagnara, 1974) – Northwest Mexico leopard frog
Lithobates megapoda (Taylor, 1942) – big-footed leopard frog
Lithobates miadis (Barbour and Loveridge, 1929) – island leopard frog
Lithobates montezumae (Baird, 1854) – Montezuma leopard frog
Lithobates neovolcanicus (Hillis and Frost, 1985) – transverse volcanic leopard frog
Lithobates okaloosae (Moler, 1985) – Florida bog frog
Lithobates omiltemanus (Günther, 1900) – Guerreran leopard frog
Lithobates onca (Cope, 1875) – relict leopard frog
Lithobates palmipes (Spix, 1824) – Amazon River frog
Lithobates palustris (LeConte, 1825) – pickerel frog
Lithobates pipiens (Schreber, 1782) – northern leopard frog
Lithobates psilonota (Webb, 2001) – smooth-backed frog
Lithobates pueblae (Zweifel, 1955) – Puebla frog
Lithobates pustulosus (Boulenger, 1883) – Mexican cascades frog
Lithobates septentrionalis (Baird, 1854) – mink frog
Lithobates sevosus (Goin and Netting, 1940) – Mississippi gopher frog
Lithobates sierramadrensis (Taylor, 1939) – Sierra Madre frog
Lithobates spectabilis (Hillis and Frost, 1985) – showy leopard frog
Lithobates sphenocephalus (Cope, 1886) – southern leopard frog
Lithobates sylvaticus (LeConte, 1825) – wood frog
Lithobates tarahumarae (Boulenger, 1917) – Tarahumara frog
Lithobates taylori (Smith, 1959) – Peralta frog
Lithobates tlaloci (Hillis and Frost, 1985) – Tlaloc's leopard frog
Lithobates vaillanti (Brocchi, 1877) – Vaillant's frog
Lithobates vibicarius (Cope, 1894) – green-eyed frog
Lithobates virgatipes (Cope, 1891) – carpenter frog
Lithobates warszewitschii (Schmidt, 1857) – Warszewitsch's frog
Lithobates yavapaiensis (Platz and Frost, 1984) – lowland leopard frog
Lithobates zweifeli (Hillis, Frost, and Webb, 1984) – Zweifel's frog

Alternatively, if Lithobates is treated as a subgenus (neotropical true frogs), then this narrower definition would contain the following species:

Rana (Lithobates) bwana Hillis and de Sá, 1988 – Rio Chipillico frog
Rana (Lithobates) juliani Hillis and de Sá, 1988 – Maya Mountains frog
Rana (Lithobates) maculata Brocchi, 1877 – highland frog
Rana (Lithobates) palmipes Spix, 1824 – Amazon River frog
Rana (Lithobates) vaillanti Brocchi, 1877 – Vaillant's frog
Rana (Lithobates) vibicaria (Cope, 1894) – green-eyed frog
Rana (Lithobates) warszewitschii Schmidt, 1857 – Warszewitsch's frog

Fossil species

The following fossil species are known, all assignable to the L. pipiens (leopard frog) complex:^[14]

†Lithobates bucella (Holman, 1965) (Early Miocene of Florida)
†Lithobates dubitus (Taylor, 1942) (Pliocene/early Pleistocene of Kansas)
†Lithobates fayeae (Taylor, 1942) (Pliocene/early Pleistocene of Kansas)
†Lithobates moorei (Taylor, 1942) (Pliocene/early Pleistocene of Kansas)
†Lithobates miocenicus (Holman, 1965) (Early Miocene of Florida)
†Lithobates robustocondylus (Taylor, 1942) (Pliocene/early Pleistocene of Kansas)
†Lithobates rexroadensis (Taylor, 1942) (Pliocene/early Pleistocene of Kansas)
†Lithobates parvissimus (Taylor, 1942) (Pliocene/early Pleistocene of Kansas)

The species described in 1942 were previously placed in their own genus, Anchylorana.^[14]

References

^ ^a ^b ^c Frost, Darrel R. (2016). "Lithobates Fitzinger, 1843". Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. Retrieved 7 August 2016.
^ Dodd, C. Kenneth (2013). Frogs of the United States and Canada. Vol. 1. The Johns Hopkins University Press. p. 20. ISBN 978-1-4214-0633-6.
^ βάτης. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project one that treads or covers
^ Hillis, David M.; Wilcox, Thomas P. (2005). "Phylogeny of the New World true frogs (Rana)" (PDF). Mol. Phylogenet. Evol. 34 (2): 299–314. doi:10.1016/j.ympev.2004.10.007. PMID 15619443. Archived from the original (PDF) on 2008-05-28.
^ Hillis, David M (2007). "Constraints in naming parts of the Tree of Life". Mol. Phylogenet. Evol. 42 (2): 331–338. doi:10.1016/j.ympev.2006.08.001. PMID 16997582.
^ ^a ^b ^c Yuan, Zhi-Yong; Zhou, Wei-Wei; Chen, Xin; Poyarkov, Nikolay A.; Chen, Hong-Man; Jang-Liaw, Nian-Hong; Chou, Wen-Hao; Matzke, Nicholas J.; Iizuka, Koji; Min, Mi-Sook; Kuzmin, Sergius L.; Zhang, Ya-Ping; Cannatella, David C.; Hillis, David M.; Che, Jing (2016). "Spatiotemporal diversification of the true frogs (genus Rana): A historical framework for a widely studied group of model organisms". Systematic Biology. 65 (5): 824–842. doi:10.1093/sysbio/syw055. hdl:2292/43460. PMID 27288482.
^ Frost, Darrel R. (2006): Amphibian Species of the World Version 3 - Petropedetidae Noble, 1931. American Museum of Natural History, New York, USA. Retrieved 2006-AUG-05., Frost, Darrel R. et al. (2006): The amphibian tree of life. Bulletin of the American Museum of Natural History. Number 297. New York.
^ Crother, B.I. (ed.) (2008): Scientific and standard English names of amphibians and reptiles of North American north of Mexico - "Standard and Common Names". Archived from the original on 2013-04-16. Retrieved 2012-04-28.. SSAR Herptological Circular 37:1-84.
^ Stuart, Bryan L. (2008): The phylogenetic problem of Huia (Amphibia: Ranidae). Mol. Phylogenet. Evol. 46(1): 49-60 doi:10.1016/j.ympev.2007.09.016 (HTMl abstract)
^ Pauly, Greg B.; Hillis, David M.; Cannatella, David C. (2009). "Taxonomic freedom and the role of official lists of species names" (PDF). Herpetologica. 65 (2): 115–128. doi:10.1655/08-031r1.1. S2CID 283839. Archived from the original (PDF) on 2011-07-25.
^ AmphibiaWeb: Information on amphibian biology and conservation. [web application]. 2012. Berkeley, California: Rana
^ "The IUCN Red List of Threatened Species. Version 2016-1". International Union for Conservation of Nature. Retrieved 7 August 2016.
^ Crother, Brian I., ed. (August 2012). Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, with comments regarding confidence in our understanding. Seventh edition. SSAR Herpetological circular No. 39 (PDF). Shoreview, MN: Society for the Study of Amphibians and Reptiles (SSAR). pp. 1–92. ISBN 978-0-916984-85-4.
^ ^a ^b ^c Sanchiz, B. (2012-01-01). "Nomenclatural notes on living and fossil amphibians". Graellsia.