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Nanorana Günther, 1896

Class: Amphibia > Order: Anura > Family: Dicroglossidae > Subfamily: Dicroglossinae > Genus: Nanorana
28 species

Nanorana Günther, 1896, Annu. Mus. Zool. Acad. Imp. Sci. St. Petersbourg, 1: 206. Type species: Nanorana pleskei Günther, 1896, by monotypy.

NannoranaWerner, 1903, Abh. Math. Physik. Cl. Bayer. Akad. Wiss., 22: 369. Incorrect subsequent spelling.

Montorana Vogt, 1924, Zool. Anz., 60: 340. Type species: Montorana ahli Vogt, 1924, by monotypy. Synonymy by Stejneger, 1927, J. Washington Acad. Sci., 17: 319.

Altirana Stejneger, 1927, J. Washington Acad. Sci., 17: 318. Type species: Altirana parkeri Stejneger, 1927, by monotypy. Regarded by Boulenger, 1920, Rec. Indian Mus., 20: 107 (and subsequently Dubois, 1981, Monit. Zool. Ital., N.S., Suppl., 15: 234), as a subgenus of Rana and close to the subgenus Paa of Rana. Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 322, considered Nanorana a subgenus of Altirana. Lü and Yang, 1995, Asiat. Herpetol. Res., 6: 69-72, and Lü and Yang, 1995, Asiat. Herpetol. Res., 6: 73-77, considered Nanorana paraphyletic with respect to Altirana and synonymized the genera. Considered a distinct genus by Zhao and Adler, 1993, Herpetol. China: 137.

Chaparana Bourret, 1939, Annexe Bull. Gen. Instr. Publique, Hanoi, 1939: 31. Type species: Rana (Chaparana) fansipani Bourret, 1939, by monotypy. Synonymy with Nanorana by Chen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 239, and Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, and Green, 2006, Bull. Am. Mus. Nat. Hist., 297: 88. Recognized as a subgenus of Nanorana by Che, Zhou, Hu, Papenfuss, Wake, and Zhang, 2010, Proc. Natl. Acad. Sci. USA, Suppl. Inform., doi:10.1073/pnas.1008415107/-/DCSupplemental: 2.

Paa Dubois, 1975, Bull. Mus. Natl. Hist. Nat. Paris, Ser. 3, Zool., 324: 1093. Type species: Rana liebigii Günther, 1860, by original designation (named originally as a subgenus of Rana). Synonymy with Nanorana by Chen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 239, and by implication of Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, and Green, 2006, Bull. Am. Mus. Nat. Hist., 297: 138. Recognized as a subgenus of Nanorana by Che, Zhou, Hu, Papenfuss, Wake, and Zhang, 2010, Proc. Natl. Acad. Sci. USA, Suppl. Inform., doi:10.1073/pnas.1008415107/-/DCSupplemental: 2.

Unculuana Fei, Ye, and Huang, 1990, Key to Chinese Amph.: 154. Type species: Rana unculuana Liu, Hu, and Yang, 1960, by original designation. Proposed as a subgenus of Paa. Synonymy with Nanorana by implication of Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, and Green, 2006, Bull. Am. Mus. Nat. Hist., 297: 318.

Quadrana Fei, Ye, and Huang, 1990, Key to Chinese Amph.: 154. Type species: Rana quadranus Liu, Hu, and Yang, 1960, by original designation. Proposed as a subgenus of Paa. Proccupied by Quadrana Caldwell and Martorell, 1952.

PaaDubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 319. Elevation from subgenus of Rana (at which rank it had been originally proposed) to generic rank.

Gynandropaa Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 319. Type species: Rana yunnanensis Anderson, 1878, by original designation. Proposed as a subgenus of Paa. Synonymy with Nanorana by implication of Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, and Green, 2006, Bull. Am. Mus. Nat. Hist., 297: 318. See comment.

Feirana Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 318. Replacement name for Quadrana Fei, Ye, and Huang, 1990. Treatment as a subgenus of Chaparana. Synonymy with Nanorana by implication of Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, and Green, 2006, Bull. Am. Mus. Nat. Hist., 297: 138. See comment.

ChaparanaDubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 318. Generic status.

FeiranaWang, Jiang, Chen, Xie, and Zheng, 2007, Acta Zootaxon. Sinica, 32: 139-145. Wang, Jiang, Xie, Chen, Dubois, Liang, and Wagner, 2009, Zool. Sci., Tokyo, 26: 500-509. Consideration as a genus.

Maculopaa Fei, Ye, and Jiang, 2010, Herpetol. Sinica, 12: 24. Type species: Rana maculosa Liu, Hu, and Yang, 1960. See comment under Dicroglossidae.

English Names

Yunnan Slow Frogs (Nanorana: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 102).

High Himalaya Frogs (Altirana [no longer recognized]: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 97).

Paa Frogs (Paa [now a subgenus]: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 102).

Asian Frogs (Chaparana [now a subgenus]: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 98).

Slow Frogs (Nanorana: Li, Zhao, and Dong, 2010, Amph. Rept. Tibet: 45).

Distribution

Himalayan region of northern Pakistan and northern India, Nepal, and western China through Myanmar, Thailand, Laos, and northern Vietnam (Lao Cai, Back Kan, Son La, Vinh Phuc, Hao Binh, and Thanh Hoa provinces) to montane southern China.

Comment

Formerly placed in the subfamily Raninae, tribe Ranini of Dubois, 1987 "1986", Alytes, 5: 39, and Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61:322. Roelants, Jiang, and Bossuyt, 2004, Mol. Phylogenet. Evol., 31: 734, suggested on the basis of molecular data that Nanorana is imbedded within Paa (Paini) and that this unit is a member of Dicroglossinae, not Raninae, as did Jiang, Dubois, Ohler, Tillier, Chen, Xie, and Stöck, 2005, Zool. Sci., Tokyo, 22: 353-362, who provided molecular evidence for paraphyly of Paa, with respect to Nanorana (including Altirana) and the polyphyly of Chaparana. Chen, Liu, Jiang, Xie, and Zheng, 2005, Herpetol. Sinica, 10: 47-51, reported on the systematics of the original three species of Nanorana. Chen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 239, placed Chaparana, Paa, and Nanorana into Nanorana on the basis that Paa is paraphyletic with respect to Nanorana and Chaparana, and this was followed by Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, and Green, 2006, Bull. Am. Mus. Nat. Hist., 297: 138, who placed Chaparana and Paa (excluding Annandia, Eripaa, Ombrana, and Quasipaa) into the synonymy of Nanorana to resolve the paraphyly of Paa with respect to Nanorana (sensu stricto). Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 313, delimited subgenera and species groups of Paa, most of which have subsequently been discarded for reasons of nonmonophyly. Dubois, 1975, Bull. Mus. Natl. Hist. Nat. Paris, Ser. 3, Zool., 324: 1111, discussed possible phylogenetic relationships of the group. Dubois, 1976, Cah. Nepal., Doc., 6, reviewed the species of Nepal and discussed the group (Paa as a subgenus of Rana). Dubois, 1987 "1986", Alytes, 5: 43, gave a summary of species groups within Paa (now in Annandia, Nanorana, Ombrana, and Quasipaa). See comments under Quasipaa and Platymantis (Ceratobatrachidae). See Tan and Wu, 1987, Acta Herpetol. Sinica, Chengdu, N.S.,, 6 (4): 35-38, for discussion of karyotypes within the group. Ohler and Dubois, 2006, Zoosystema, 28: 769-784, presented a taxonomy that is difficult to address because it did not take into account the molecular results of Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, and Green, 2006, Bull. Am. Mus. Nat. Hist., 297, and in which Ohler and Dubois explicitly recognized paraphyletic (genus Chaparana and its subgenus Paa) and polyphyletic (subgenus Chaparana) taxa, contrary to standard taxonomic practice and their own phylogenetic evidence. Rather than adopt an explicitly misleading taxonomy, I (DRF) retain Ohler and Dubois' Chaparana within (and as a synonym of) a monophyletic Nanorana, but reflect their nomenclature in the synonymies. Where their conclusions are not rejected by their own or other's (e.g., Frost et al., 2006) phylogenetic evidence, I have adopted their taxonomic novelties (e.g., recognition of Chrysopaa and Allopaa), although this is an interim measure until such time as an adequate analysis is completed and certainly does not reflect any equivalency of these taxa (as suggested by Wang, Jiang, Xie, Chen, Dubois, Liang, and Wagner, 2009, Zool. Sci., Tokyo, 26: 500-509). It is unclear what the results will be when the morphology and molecular evidence are combined, but it clearly will not support the taxonomy suggested by Ohler and Dubois (2006). Without discussion the junior synonym Annandia was treated as a distinct genus within Limnonectini by Dubois, 2005, Alytes, 23: 16; previously Annandia had been considered a subgenus of Chaparana (= Nanorana). Jiang, Dubois, Ohler, Tillier, Chen, Xie, and Stöck, 2005, Zool. Sci., Tokyo, 22: 353-362, did not mention Annandia in their revision of Paini; as a result Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, and Green, 2006, Bull. Am. Mus. Nat. Hist., 297: 241, followed Dubois, 2005, in recognizing Annandia. Ohler and Dubois, 2006, Zoosystema, 28: 769-784, on the basis of a parsimony analysis of morphology suggested that Annandia is more closely related to Limnonectes and Euphlyctis than to former members of Paini. Wiens, Sukumaran, Pyron, and Brown, 2009, Evolution, 63: 1217-1231, corroborated the monophyly of Nanorana sensu Frost et al., 2006. Wang, Jiang, Xie, Chen, Dubois, Liang, and Wagner, 2009, Zool. Sci., Tokyo, 26: 500-509, provided a small phylogenetic study with sparse taxon sampling that allowed the authors to recognize Feirana as a genus. However, their analysis, employed only one terminal of Nanorana so the hypothesis that recognition of Feirana would result in the paraphyly of Nanorana has not been rejected and the synonymy of Feirana is here retained pending the evidentiary resolution of this problem. Recognized as a subgenus of Nanorana by Che, Zhou, Hu, Papenfuss, Wake, and Zhang, 2010, Proc. Natl. Acad. Sci. USA, Suppl. Inform., doi:10.1073/pnas.1008415107/-/DCSupplemental: 2-3. Yang, Wang, Hu, and Jiang, 2011, Asian Herpetol. Res., Ser. 2, 2: 72-86, suggested Feirana as a genus, but restricted it in a way that would render Nanorana paraphyletic (given the tree of Che, Zhou, Hu, Papenfuss, Wake, and Zhang, 2010, Proc. Natl. Acad. Sci. USA, 107: 13765-13770). For this reason, Feirana is retained within the synonymy of Nanorana for purposes of this catalogue. Although they employed an antiquated nonmonophyletic taxonomy which makes appreciating their results difficult, Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543-583, in their study of Genbank sequences found Nanorana to be monophyletic and the sister taxon of Quasipaa (but see comments under Chrysopaa , Ombrana, and Allopaa), as well as providing a tree of exemplar species, although explicitly embracing an out-dated taxonomy (e.g., including the non-monophyletic Chaparana and Paa) that is inconsistent with their own tree topology when the current taxonomy (the one employed here), which is consistent with their recovered tree.

Contained taxa (28 sp.):

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