Northern red-legged frog Rana aurora, copyright John Battaso.

Belongs within: Ranidae.

Rana is a diverse genus of frogs with smooth skin, strong hindlimbs and webbed toes.

Rana, Rana, Rana
Published 26 June 2007

Rana is a large primarily Holarctic genus of frogs, and probably the inspiration for most depictions of frogs in the world. Well-known species are the edible frog Rana × esculenta (actually not a true species but a hybrid) and the European common frog Rana temporaria.

Edible frog Rana esculenta, copyright Grand-Duc, Niabot.

I thought I’d look up the info on Rana over lunchtime. Pretty soon, my head was swimming. The genus Rana has been hit with two major investigations in recent years, both of which have received some frosty responses. Frost et al. (2006) in their investigation of the ‘Amphibian Tree of Life’ divided Rana between more than fifteen smaller genera to remove its previous paraphyly (for instance, the above-mentioned Rana esculenta would become Pelophylax esculentus). As happens with any wholesale name change, there has been quite a bit of outcry at the idea of having to update the filing systems. Also, a number of authors have felt that the number of taxa sampled by Frost et al. was not enough to inspire confidence in their results. The review by Wiens (2007) was particularly vitriolic—the scientific equivalent of attempting to hold the subject down and kick them repeatedly in the teeth. Smith and Chiszar (2006) have suggested the more mollifying approach of treating Frost et al.‘s various genera as subgenera, though unless one was willing to accept a paraphyletic genus this would also require sinking some well-established genera such as Staurois within Rana. Division of the genus Rana was also supported by Che et al. (2007).

The other cause of debate was perpetrated by Hillis & Wilcox (2005), who investigated the phylogeny of New World species of ‘Rana’ (most of which would belong to Lithobates in the Frost et al. system). The problem came when Hillis & Wilcox suggested a whole series of subgeneric taxa for nested groups of species that they defined according to the rules of the PhyloCode, but also allowed for use under the ICZN as subgenera. Debate promptly exploded about whether Hillis & Wilcox’s names were validly published and usable (Dubois, 2006, 2007; Hillis, 2007). Compared to this argument, Frost et al.‘s division appears quite simple. I may return to this in a later post, if my brain doesn’t implode first. Check out the Dubois (2006) paper in particular—Dubois thinks that the answer to our problems is to make the ICZN more complicated. No. Thank. You.

Systematics of Rana
Rana Linnaeus 1758FG06 (see below for synonymy)
|--R. weiningensis Liu, Hu & Yang 1962PW11, FG06 [=R. (Pseudorana) weiningensisFG06]
`--+--+--R. shuchinae Liu 1950PW11, FG06 [=Pelophylax shuchinaeFG06]
| `--+--+--+--R. luteiventris Thompson 1913PW11, FG06
| | | `--R. pretiosa Baird & Girard 1853PW11, FG06 [=R. (Aurorana) pretiosaFG06, R. temporaria pretiosaW06]
| | `--+--R. boylii Baird 1854PW11, FG06 [=R. (Amerana) boyliiFG06]
| | `--+--R. aurora Baird & Girard 1852PW11, FG06 [=R. (Aurorana) auroraFG06]
| | `--+--R. cascadae Slater 1939 [=R. (Aurorana) cascadae]FG06
| | `--R. muscosa Camp 1917FG06 (see below for synonymy)
| `--+--R. tagoi Okada 1928PW11, FG06
| `--+--+--R. johnsi Smith 1921PW11, FG06 [=R. (Pseudorana) johnsiFG06]
| | `--R. zhengi Zhao 1999PW11, FG06
| `--+--+--+--R. amurensis Boulenger 1886FB17, FG06 [=Rana temporaria amurensisL50]
| | | `--R. kunyuensis Lu & Li 2002PW11, FG06
| | `--+--R. arvalis Nilsson 1842PW11, FG06 [incl. R. oxyrrhinus Steenstrup 1846M01]
| | | |--R. a. arvalisK08
| | | `--R. a. wolterstorffiK08
| | `--+--R. japonica Boulenger 1879FB17, FG06 (see below for synonymy)
| | `--+--R. chaochiaoensis Liu 1946PW11, FG06
| | `--+--R. omeimontis Ye & Fei 1993PW11, FG06
| | `--+--R. longicrus Stejneger 1898PW11, FG06
| | `--R. zhenhaiensis Ye, Fei & Matsui 1995PW11, FG06
| `--+--+--R. ornativentris Werner 1903PW11, FG06
| | `--+--+--R. dybowskii Günther 1876PW11, FG06
| | | `--R. pirica Matsui 1991PW11, FG06
| | `--+--R. huanrenensis Fei, Ye & Huang 1991PW11, FG06
| | `--+--R. chensinensis David 1875FB17, FG06 (see below for synonymy)
| | `--R. kukunoris Nikolskii 1918PW11, FG06
| `--+--+--R. asiatica Bedriaga 1898PW11, FG06 [=R. temporaria var. asiaticaL50]
| | `--R. sauteri Boulenger 1909FG06 [=*Pseudoamolops sauteriFG06, PW11, Pseudorana sauteriFG06]
| `--+--+--R. latastei Boulenger 1879PW11, FG06
| | `--+--R. dalmatina Fitzinger in Bonaparte 1839PW11, FG06
| | `--R. graeca Boulenger 1891PW11, FG06
| `--+--+--R. holsti Boulenger 1892PW11, FG06 [=Babina holstiFG06]
| | `--R. macrocnemis Boulenger 1885PW11, FG06
| | |--R. m. macrocnemisRR00
| | `--R. m. angeloi Bogachev 1927 (n. d.) [incl. R. kisatibensis Riabinin 1928]RR00
| `--+--+--R. okinavana Boettger 1895PW11, FG06
| | `--R. tsushimensis Stejneger 1907PW11, FG06
| `--+--+--*R. temporaria Linnaeus 1758FG06, FB17, FG06
| | `--R. pyrenaica Serra-Cobo 1993PW11, FG06
| `--+--R. iberica Boulenger 1879PW11, FG06
| `--R. italica Dubois 1987PW11, FG06
`--+--+--R. sylvatica Le Conte 1825PW11, FG06 [=Lithobates sylvaticusFG06]
| `--+--R. catesbeiana Shaw 1802PW11, FG06 (see below for synonymy)
| `--+--+--R. grylio Stejneger 1901PW11, FG06 [=Lithobates grylioFG06, R. (Aquarana) grylioFG06]
| | `--R. septentrionalis Baird 1854PW11, FG06 [=Lithobates septentrionalisFG06]
| `--+--R. virgatipes Cope 1891PW11, FG06 [=Lithobates virgatipesFG06, R. (Aquarana) virgatipesFG06]
| `--+--R. hecksheri Wright 1924PW11, FG06 [=Lithobates hecksheriFG06, R. (Aquarana) heckscheriFG06]
| `--+--R. clamitans Latreille 1801PW11, FG06 [=Lithobates clamitansFG06, R. (Aquarana) clamitansFG06]
| `--R. okaloosae Moler 1985 [=Lithobates okaloosae]FG06
`--+--+--+--R. sierramadrensis Taylor 1939PW11, FG06 [=Lithobates sierramadrensisFG06, R. (Sierrana) sierramadrensisFG06]
| | `--+--+--R. psilonota Webb 2001PW11, FG06 [=Lithobates psilonotaFG06]
| | | `--R. zweifeli Hillis, Frost & Webb 1984 [=Lithobates zweifeli, R. (Zweifelia) zweifeli]FG06
| | `--+--R. pustulosa Boulenger 1883FG06 (see below for synonymy)
| | `--R. tarahumarae Boulenger 1917 [=Lithobates tarahumarae, R. (Zweifelia) tarahumarae]FG06
| `--+--+--R. maculata Brocchi 1877PW11, FG06 [=Lithobates maculatusFG06, R. (Sierrana) maculataFG06]
| | `--+--R. vibicaria Cope 1894 [=Lithobates vibicarius, R. (Trypheropsis) vibicaria]FG06
| | `--R. warszewitschii Schmidt 1857 [=Lithobates warszewitschii, R. (Trypheropsis) warszewitschii]FG06
| `--+--+--R. bwana Hillis & de Sá 1988PW11, FG06 [=Lithobates bwanaFG06]
| | `--R. palmipes Spix 1824 [=R. (Lithobates) palmipes]FG06
| `--+--R. juliani Hillis & de Sá 1988 [=Lithobates juliani]FG06
| `--R. vaillanti Brocchi 1877 [=R. (Lithobates) vaillanti]FG06
`--+--+--+--+--R. areolata Baird & Girard 1852PW11, FG06 [=Lithobates areolatusFG06, R. (Pantherana) areolataFG06]
| | | `--R. palustris LeConte 1825PW11, FG06 [=Lithobates palustrisFG06, R. (Pantherana) palustrisFG06]
| | `--+--R. capito LeConte 1855PW11, FG06 [=Lithobates capitoFG06, R. (Pantherana) capitoFG06]
| | `--R. sevosa Goin & Netting 1940 [=Lithobates sevosus]FG06
| `--+--R. pipiens Schreber 1782PW11, FG06 [=Lithobates pipiensFG06, R. (Pantherana) pipiensFG06]
| `--+--+--R. chiricahuensis Platz & Mecham 1979 (see below for synonymy)FG06
| | `--R. subaquavocalisFG06
| `--+--R. dunni Zweifel 1957 [=Lithobates dunni]FG06
| `--+--R. megapoda Taylor 1942HFW83, FG06 [=Lithobates megapodaFG06]
| `--R. montezumae Baird 1854 [=Lithobates montezumae, R. (Pantherana) montezumae]FG06
`--+--+--R. magnaocularis Frost & Bagnara 1974PW11, FG06 (see below for synonymy)
| `--+--R. onca Cope 1875 [=Lithobates onca]FG06
| `--R. yavapaiensis Platz & Frost 1984 [=Lithobates yavapaiensis, R. (Pantherana) yavapaiensis]FG06
`--+--+--R. sphenocephala Cope 1886PW11, FG06 (see below for synonymy)
| `--+--R. blairi Mecham, Littlejohn et al. 1973 [=Lithobates blairi, R. (Pantherana) blairi]FG06
| `--+--R. berlandieri Baird 1859 [=Lithobates berlandieri, R. (Pantherana) berlandieri]FG06
| `--+--R. neovolcanica Hillis & Frost 1985 [=Lithobates neovolcanicus]FG06
| `--R. tlaloci Hillis & Frost 1985 [=Lithobates tlaloci]FG06
`--+--+--R. forreri Boulenger 1883PW11, FG06 [=Lithobates forreriFG06, R. (Pantherana) forreriFG06]
| `--R. omiltemana Günther 1900PW11, FG06 [=Lithobates omiltemanusFG06]
`--+--R. spectabilis Hillis & Frost 1985PW11, FG06 (see below for synonymy)
`--+--R. brownorum Sanders 1973PW11, FG06 [=Lithobates brownorumFG06]
`--+--R. macroglossa Brocchi 1877PW11, FG06 [=Lithobates macroglossaFG06]
`--R. taylori Smith 1959 [=Lithobates taylori]FG06

Rana incertae sedis:
R. agilisW06
R. albotuberculata Inger 1954 [=Hydrophylax albotuberculatus]FG06
R. amnicola Perret 1977 [=Hydrophylax amnicola]FG06
R. anlungensis Liu & Hu 1973 [=Huia anlungensis, R. (Odorrana) anlungensis]FG06
R. antiquaRR00
R. aquensisRR00
R. arborea Linnaeus 1758L58
R. asperrima Perret 1977 [=Hydrophylax asperrima]FG06
R. attigua Inger, Orlov & Darevsky 1999 [=Sylvirana attigua]FG06
R. aurata Günther 2003 [=Sylvirana aurata]FG06
R. bannanica Rao & Yang 1997 [=Sylvirana bannanica]FG06
R. basaltica Young 1936RR00
R. boans Linnaeus 1758L58
R. boulengeri Günther 1889 [incl. R. tibetana Boulenger 1917]L50
R. caldwelli Schmidt 1925 [=Babina caldwelli, Nidirana caldwelli]FG06
R. camerani Boulenger 1886FG06
R. cantabrigensisW06
R. celebensis Peters 1872 [=Sylvirana celebensis]FG06
R. chevronta Hu & Ye 1978FG06
R. chichicuahutla Cuellar, Méndez-De La Cruz & Villagrán-Santa Cruz 1996 [=Lithobates chichicuahutla]FG06
R. chinensis Osbeck 1771 (n. d.)L50, M01 [=R. esculenta var. chinensisM01]
R. chitwanensis Das 1998 [=Sylvirana chitwanensis]FG06
R. chosenica Okada 1931 [=Pelophylax chosenicus]FG06
R. christyiPB27
R. clamataJ23
R. cornuta Linnaeus 1758L58
R. crassiovis Boulenger 1920 [=Hydrophylax crassiovis]FG06
R. crassipesKF69
R. danieli Pillai & Chanda 1977 [=Sylvirana danieli]FG06
R. danubianaRR00
R. darlingi Boulenger 1902 [=Hydrophylax darlingi]FG06
R. daunchina Chang 1933 [=Babina daunchina]FG06
R. debussyi van Kampen 1910 [=Pulchrana debussyi]FG06
R. demarchii Scortecci 1929 [=Pelophylax demarchii]FG06
R. draytonii Baird & Girard 1852FG06 [=R. aurora draytoniiG86]
R. elberti Roux 1911 [=Sylvirana elberti]FG06
R. ephippium (n. d.)RR00
R. everetti Boulenger 1882 [=Hydrophylax everetti]FG06
R. exiliversabilis Li, Ye & Fei 2001 [=Odorrana (Bamburana) exiliversabilis, Huia exiliversabilis]FG06
R. feae Boulenger 1887L50
R. feyae Taylor 1942RR00
R. fisheri Stejneger 1893FG06 [=Rana onca fisheriD81, R. pipiens fisheriH81, Lithobates fisheriFG06]
R. florensis Boulenger 1897 [=Sylvirana florensis]FG06
R. fusca Rösel 1758 (see below for synonymy)M01
R. garoensis Boulenger 1920 [=Sylvirana garoensis]FG06
R. garritor Menzies 1987 [=Sylvirana garritor]FG06
R. gibbosa Linnaeus 1758L58
R. grandocula Taylor 1920 [=Pulchrana grandocula]FG06
R. greeniiWKW73
R. grisea van Kampen 1913 [=Sylvirana grisea]FG06
R. hainanensis Fei, Ye & Li 2001 [=Huia hainanensis, R. (Odorrana) hainanensis]FG06
R. halecinaJ23
R. hipparionum Schlosser 1924RR00
R. holtzi Werner 1898FG06
R. hyla Linnaeus 1758L58
R. jaegeriRR00
R. jerboaKF69
R. johni Blair 1965 [=Lithobates johni]FG06
R. johnsi Smith 1921FG06
R. johnsoni LaRivers 1953RR00
R. kampeni Boulenger 1920 [=Hydrophylax kampeni]FG06
R. kreffti Boulenger 1882 [=Sylvirana kreffti]FG06
R. kuhlii Duméril & Bibron 1841 [incl. Nyctibatrachus sinensis Peters 1882]L50
R. lemairei de Witte 1921 [=Hydrophylax lemairei]FG06
R. lemosespinali Smith & Chiszar 2003 [=Lithobates lemosespinali]FG06
R. leptoglossa Cope 1868 [=Sylvirana leptoglossa]FG06
R. limnocharis Boie in Wiegmann 1835L50
R. lini Chou 1999 [=Babina lini]FG06
R. longipes Perret 1960 [=Hydrophylax longipes]FG06
R. luschitzanaRR00
R. macrodonM89
R. macrops Boulenger 1897 [=Hydrophylax macrops]FG06
R. mangyanum Brown & Guttman 2002 [=Pulchrana mangyanum]FG06
R. margariana Anderson 1879 [=Sylvirana margariana]FG06
R. marginata Linnaeus 1758L58
R. marina Linnaeus 1758L58
R. mascareniensisPB27
R. meadensis (n. d.)RR00
R. megapodaC78
R. mehelyiJ63
R. melanomenta Taylor 1920 [=Pulchrana melanomenta]FG06
R. meriani Meyer 1853RR00
R. miadis Barbour & Loveridge 1929 [=Lithobates miadis]FG06
R. miocenica Holman 1965 [incl. Leptodactylus abavus Holman 1965, R. abavus]RR00
R. modestaB28
R. moellendorffi Boettger 1893 [=Pulchrana moellendorffi]FG06
R. moluccana Boettger 1895 [=Sylvirana moluccana]FG06
R. montivaga Smith 1921 [=Sylvirana montivaga]FG06
R. moorei (Taylor 1942)C78, RR00 [=Anchylorana mooreiRR00]
R. mortenseni Boulenger 1903 [=Sylvirana mortenseni]FG06
R. multidenticulata Chou & Lin 1997FG06
R. namiyei Stejneger 1901I92
R. nasuta Li, Ye & Fei 2001 [=Huia nasuta, Odorrana (Bamburana) nasuta]FG06
R. nigrolineata Liu & Hu 1960 [=Pelophylax nigrolineatus]FG06
R. nigrotympanica Dubois 1992 [=Sylvirana nigrotympanica]FG06
R. noeggerathi Meyer 1852RR00
R. novaeguineae van Kampen 1909 [=Sylvirana novaeguineae]FG06
R. occidentalis Perret 1960 [=Hydrophylax occidentalis]FG06
R. occipitalisWKW73
R. ocellata Linnaeus 1758L58
R. ornatissimaPB27
R. oxyrhynchusPB27
R. papua Lesson 1831 [=Sylvirana papua]FG06
R. paradoxa Linnaeus 1758L58
R. parkeriana Mertens 1938 [=Hydrophylax parkerianus]FG06
R. persimilis van Kampen 1923 [=Sylvirana persimilis]FG06
R. phrynoides Boulenger 1917L50
R. pipa Linnaeus 1758L58
R. plicata (Filhol 1876)RR00
R. pliocenica Zweifel 1964RR00
R. psaltes Kuramoto 1985 [=Babina psaltes]FG06
R. pueblae Zweifel 1955 [=Lithobates pueblae]FG06
R. pueyoiRR00
R. quellenbergiRR00
R. raraRR00
R. rexroadensis Taylor 1942RR00
R. rubeta Linnaeus 1758L58
R. rugataB28
R. sakuraii Matsui & Matsui 1990FG06
R. salzhausenensisRR00
R. sangzhiensis Shen 1986 [=R. (Pseudorana) sangzhiensis]FG06
R. sieblosensisRR00
R. similis Günther 1873 [=Pulchrana similis]FG06
‘Ranomorphus similis’ Ratnikov 1993 non Rana similis Günther 1873RR00
R. spinosaL50
R. straussi Špinar 1980RR00
R. subaspera Barbour 1908 [=Babina subaspera]FG06
R. supragrisea Menzies 1987 [=Sylvirana supragrisea]FG06
R. tenggerensis Zhao, Macey & Papenfuss 1988 [=Pelophylax tenggerensis]FG06
R. terentievi Mezhzherin 1992 [=Pelophylax terentievi]FG06
R. tigerinaL50
|--R. t. tigerinaL50
`--R. t. rugulosa Wiegmann 1835L50
R. tipanan Brown, McGuire & Diesmos 2000 [=Hydrophylax tipanan]FG06
R. troscheliRR00
R. typhonia Linnaeus 1758L58
R. tytleri Theobald 1868 [=Hylarana tytleri]FG06
R. utriculariaPU81
R. valida (n. d.)RR00
R. varians Boulenger 1894FG06 [=Sanguirana variansFG06, R. papua variansB28]
R. variegata Linnaeus 1758L58
R. ventricosa Linnaeus 1758L58
R. virescensW06
R. vittigeraL50
R. volhynicaRR00
R. volkerjane Günther 2003 [=Sylvirana volkerjane]FG06
R. yalpugiensis (Ratnikov 1993) (n. d.)RR00
R. yunnanensis Anderson 1878L50

Rana Linnaeus 1758FG06 [incl. Amerana Dubois 1992FG06, Amnirana Dubois 1992FG06, Aquarana Dubois 1992FG06, Aurorana Dubois 1992FG06, Conrana Sangster 2005F05, Eburana Dubois 1992FG06, Glandirana Fei, Ye & Huang 1991FG06, Lithobates Fitzinger 1843FG06, Nidirana Dubois 1992FG06, Pantherana Dubois 1992FG06, Pseudoamolops Jiang, Fei et al. 1997FG06, Pseudorana Dubois 1992FG06, Pulchrana Dubois 1992FG06, Ranomorphus Ratnikov 1993RR00, Sierrana Dubois 1992FG06, Trypheropsis Cope 1868FG06, Tylerana Dubois 1992FG06, Zweifelia Dubois 1992FG06]

Rana catesbeiana Shaw 1802PW11, FG06 [=R. (Aquarana) catesbeianaFG06, *Conrana catesbeianaF05, Lithobates catesbeianusFG06]

Rana chensinensis David 1875FB17, FG06 [=R. temporaria chensinensisL50; incl. R. weigoldi Vogt 1924L50]

Rana chiricahuensis Platz & Mecham 1979 [=Lithobates chiricahuensis, R. (Pantherana) chiricahuensis]FG06

Rana fusca Rösel 1758 [incl. R. temporaria var. bosniensis, R. cruenta Pallas 1831, R. muta Laurenti 1768, R. platyrrhinus Steenstrup 1846]M01

Rana japonica Boulenger 1879PW11, FG06 [=R. temporaria var. japonicaL50; incl. R. silvatica Swinhoe 1870L50]

Rana magnaocularis Frost & Bagnara 1974PW11, FG06 [=Lithobates magnaocularisFG06, R. (Pantherana) magnaocularisFG06]

Rana muscosa Camp 1917FG06 [=R. (Amerana) muscosaFG06, R. boylii muscosaG86; incl. R. boylii sierraeG86]

Rana pustulosa Boulenger 1883FG06 [=Lithobates pustulosusFG06, R. (Zweifelia) pustulosaFG06; incl. R. sinaloaeG86]

Rana spectabilis Hillis & Frost 1985PW11, FG06 [=Lithobates spectabilisFG06, R. (Pantherana) spectabilisFG06]

Rana sphenocephala Cope 1886PW11, FG06 [=Lithobates sphenocephalusFG06, R. (Pantherana) pipiens sphenocephalaFG06, CM07]

*Type species of generic name indicated


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