Hylarana Tschudi, 1838

English Names

Golden-backed Frogs (Du Preez and Carruthers, 2009, Compl. Guide Frogs S. Afr.: 450).

White-lipped Frogs (Amnirana: Channing and Howell, 2006, Amph. E. Afr.: 255).

Caped Frogs (Humerana: Dinesh, Radhakrishnan, Deepak, and Kulkarni, 2023, Fauna India Checklist, vers. 5.0 : 10).

Golden-back Frogs (Insylvirana: Dinesh, Radhakrishnan, Deepak, and Kulkarni, 2023, Fauna India Checklist, vers. 5.0 : 10).

Wood Frogs (Sylvirana: Dinesh, Radhakrishnan, Deepak, and Kulkarni, 2023, Fauna India Checklist, vers. 5.0 : 11).

Distribution

Subsaharan Africa south to Namibia and Zimbabwe; subtropical and tropical Asia to throughout the Sunda Shelf and Philippines to Australia. 

Comment

 Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 1–738, found Pulchrana and Amnirana to be monophyletic, with Chalcorana to be paraphyletic with respect to these two taxa. They recovered Sylvirana, Papurana, Hydrophylax, Hylarana (sensu stricto) and Indosylvirana as polyphyetic. On this basis they place all of the genera into the synonymy of Hylarana, although this taxonomic arrangement is probably not going to last. For this reason I retain all of the comments attached to the nominal genera seperately below. 

Hylarana was considered a genus (and redelimited) most recently by Oliver, Prendini, Kraus, and Raxworthy, 2015, Mol. Phylogenet. Evol., 90: 176–192, who revised the group, recognizing Abavorana, Pulchrana, Chalcorana, Amnirana, Humerana, a much reduced Hylarana, Sylvirana, Hydrophylax, and Papurana as genera. Prior to that revision  Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1-13; Chen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 237, had recognized a much larger Hylarana out of the genera recognized by the revision of Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 248, who provided a long taxonomic history of this nominal taxon and a partial revision. Bossuyt, Brown, Hillis, Cannatella, and Milinkovitch, 2006, Syst. Biol., 55: 584, provided evidence that Hylarana (sensu Frost et al., 2006) is imbedded within a paraphyletic Sylvirana (which has now been redelimited itself). Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 324, on the basis of subjective similarity considered his subgenera Hydrophylax and Amnirana to be in a subsection Hydrophylax of his section Hylarana, and Chalcorana in the same section, but subsection Hylarana. Matsui, Shimada, Ota, and Tanaka-Ueno, 2005, Mol. Phylogenet. Evol., 37: 733-742, and Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 1-370, found this system of sections and subsections (and some subgenera) to be poorly correlated with evolutionary history. Bossuyt, Brown, Hillis, Cannatella, and Milinkovitch, 2006, Syst. Biol., 55: 584, provided evidence for the polyphyly of Hydrophylax (sensu Frost et al., 2006); in their results Hydrophylax luzonensis (now Sanguirana luzonensis) is most closely related to Sanguirana sanguinea and with Hydrophylax galamensis and Hydrophylax chalconota imbedded within another group containing Pulchrana and at least one species of Sylvirana (nicobariensis). Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1-13, placed Hydrophylax (sensu Frost et al., 2006) into the synonymy of Hylarana. See comment under Humerana, which likely renders Hylarana, in the sense of this catalogue, paraphyletic. Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 250, retained Pulchrana as a genus, pending its phylogenetic placement, and rejected the system of sections and subsections suggested by Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61. Bossuyt, Brown, Hillis, Cannatella, and Milinkovitch, 2006, Syst. Biol., 55: 584, provided evidence that Pulchrana is imbedded within a paraphyletic Sylvirana. Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1-13, provided evidence that Pulchrana is imbedded within Hydrophylax (including Chalcorana) of Frost et al. (2006) and they regarded it as a junior synonym of Hylarana. Bossuyt, Brown, Hillis, Cannatella, and Milinkovitch, 2006, Syst. Biol., 55: 584, provided evidence for the paraphyly of Sylvirana, with with respect to Pulchrana, Hylarana (e.g., Hylarana erythraea), and elements of Hydrophylax (e.g., Hydrophylax galamensis, Hydrophylax chalconota, sensu Frost et al., 2006) imbedded within it. Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 3, regarded Sylvirana as a junior synonym of Hylarana. Kraus and Allison, 2007, Herpetol. Monogr., 21: 33-75, reviewed the Hylarana papua species group (as Rana papua) in Papua New Guinea. Fei, Ye, Jiang, and Xie, 2008, Acta Zootaxon. Sinica, 33: 199-206, provided a key to the species of the Hylarana nigrovittata group in China. See comment under Rana cangyuanensis. Wiens, Sukumaran, Pyron, and Brown, 2009, Evolution, 63: 1217-1231, presented results that suggest that Hylarana (sensu Che et al., 2007) is paraphyletic with respect to Sanguirana, Glandirana, Rana, and Lithobates, suggesting that partitions of this nominal taxon should be expected. Du Preez and Carruthers, 2009, Compl. Guide Frogs S. Afr.: 450-455, provided accounts and a key for the species of southern Africa. Fei, Hu, Ye, and Huang, 2009, Fauna Sinica, Amph. 3: 1124-1196, provided keys and accounts and suggested a system of subgenera (including Babina of this catalog) and species groups not formally substantiated by synapomorphy. Kurabayashi, Yoshikawa, Sato, Hayashi, Oumi, Fujii, and Sumida, 2010, Mol. Phylogenet. Evol., 56: 543-553, suggested that Hylarana is close to Glandirana. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543-583, confirmed the likely monophyly (see comments under Humerana and Pterorana) of Hylarana although this is obscured by their adoption of an antiquated and non-monophyletic taxonomy. Biju, Garg, Mahony, Wijayathilaka, Senevirathne, and Meegaskumbura, 2014, Contr. Zool., Amsterdam, 83: 269–335, revised the delimited the Hylarana aurantiaca group (now Indosylvirana) of India, the Hylarana flavescens group (now Indosylvirana) of India, Hylarana malabarica group (now Hydrophylax) of India and Sri Lanka, and the Hylarana temporalis group (now Indosylvirana) of Sri Lanka. 

Sylvirana: Removed from the synonymy of Hylarana by Oliver, Prendini, Kraus, and Raxworthy, 2015, Mol. Phylogenet. Evol., 90: 188, where it had been placed by Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 3. See comment under Hylarana for access to relevant revisionary literature. Sheridan and Stuart, 2018, PLoS One, 13(3: e0192766): 1–48, reported on the molecular phylogenetics of the species similar to Sylvirana nigrovittata of Myanmar, Thailand, Laos, Vietnam, Cambodia, and peninsular Malaysia. Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 423, recovered Sylvirana as polyphyletic, one species, Sylvirana guentheri as the sister taxon of Humerana miopus, and another group with Hylarana latouchii imbedded within it (and including the type species, Sylvirana nigrovittata). On this basis, and the general chaotic nature of hylaranine genera, these authors place Sylvirana into the synonymy of Hylarana. 

Papurana: Removed from the synonymy of Hylarana by Oliver, Prendini, Kraus, and Raxworthy, 2015, Mol. Phylogenet. Evol., 90: 190 (who considered it the sister taxon of Hydrophylax), where it had been placed by Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 3.  See comment under Hylarana for access to relevant revisionary literature.  Donnellan, Aplin, and Bertozzi, 2010, Zootaxa, 2496: 49-62, reported on phylogenetics of the New Guinean species, and noted that Papurana is paraphyletic with respect to Tylerana. Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 423, recovered Papurana as polyphyletic in two clades, one (Papurana daemeli, Papurana papua, Papua arfaki, and Papurana jimiensis) forming the weakly resolved sister taxon of Hydrophylax leptoglossa, and another (Papurana milleti and Papurana attigua) forming the weakly resolved sister taxon of another group of Hydrophylax. Whether this is due of alignment issues (DRF's guess) or that previous work has gotten everything terribly wrong, is unclear. Regardless, these authors placed Papurana into the synonymy of Hylarana. Reilly, Arifin, Stubbs, Karin, Kaiser, Frederick, Arida, Iskandar, and McGuire, 2022, Zootaxa, 5150: 591–599, treated Papurana as a subgenus of Hylarana without comment on its monophyly in their mtDNA study of the phylogenetics of the populations in the Lesser Sundas. 

Amnirana: Removed from the synonymy of Hylarana by Oliver, Prendini, Kraus, and Raxworthy, 2015, Mol. Phylogenet. Evol., 90: 186, where it had been placed by Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 3. See comment under Hylarana for access to relevant revisionary literature. Amnirana was previous recognized as a genus by Channing, 2001, Amph. Cent. S. Afr.: 266. Synonymies of African species of "Hylarana" (now Amnirana) provided by Perret, 1977, Rev. Suisse Zool., 84: 842-844. Rödel and Bangoura, 2004, Tropical Zool., 17: 201-232, provided a key to the species of West Africa (as Amnirana). Jongsma, Barej, Barratt, Burger, Conradie, Ernst, Greenbaum, Hirschfeld, Leaché, Penner, Portik, Zassi-Boulou, Rödel, and Blackburn, 2018, Mol. Phylogenet. Evol., 120: 274–285, reported on molecular phylogenetics and biogeography in sub-Saharan Africa. Channing and Rödel, 2019, Field Guide Frogs & Other Amph. Afr.: 386–395, provided brief accounts, photographs, and range maps for the species. Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 423, found their exemplars of Amnirana (Amnirana galamensis, Amnirana albolabris, and Amnirana lepus) to form a monophyletic group. However, due to the general poor resolution of hylaranine genera, these authors placed Amnirana into the synonymy of Hylarana. 

Pulchrana: Removed from the synonymy of Hylarana by  Oliver, Prendini, Kraus, and Raxworthy, 2015, Mol. Phylogenet. Evol., 90: 188, where it had been placed by Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 3. See comment under Hylarana for access to relevant revisionary literature. Brown and Guttman, 2002, Biol. J. Linn. Soc., 76: 393-461, discussed the phylogenetics of the Rana signata complex (Pulchrana) of the Philippines and Borneo. Chan, Abraham, Grismer, and Brown, 2020, Raffles Bull. Zool., 68: 880–890, summarized the state of knowledge of the Pulchrana picturata group (Pulchrana sundabarat and Pulchrana picturata). Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 423, recovered Pulchrana as monophyletic, the sister of the bulk of their Chalcorana samples, but, presumably because of the nonmonophyly of other hylaranines, these authors placed Pulchrana into the synonymy of Hylarana. 

Indosylvirana: See comment under Hylarana for access to relevant revisionary literature. Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 423, recovered Indosylvirana as polyphyletic with two species forming a near-basal monophyletic group within the hylaranines, Indosylvirana aurantiaca + Indosylvirana temporalis, and another species, Indosylvirana nicobariensis, about midway up the spine of the hylaranine tree. Whether this is due to alignment problems in the overall analysis (DRF's guess) of real polyphyly is at this juncture unknown. Nevertheless, the authors placed all genera of hylarainines other than Abavorana into the synonymy of Hylarana on the basis of their recovery of widespread non-monophyly of genera. 

Hydrophylax: Removed from the synonymy of Hylarana by Oliver, Prendini, Kraus, and Raxworthy, 2015, Mol. Phylogenet. Evol., 90: 187, where it had been placed by Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 3. Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 423, recovered Hydrophylax as polyphyletic, with one exemplar, Hydrophylax leptoglossa being the sister of one cluster of Papurana, and Hydrophylax malabarica being the sister of another cluster of Papurana. For this reason, and a general inability to recover monophyletic hylaranine genera, the authors placed Hydrophylax into the synonymy of Hylarana. 

Humerana: In the Section Hylarana, subsection Hydrophylax, subgenus Humerana of Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 324. Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 250, regarded Humerana as a genus, rather than a subgenus of Rana, pending its phylogenetic placement. Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1–13, did not address this taxon. Matsui, Shimada, Ota, and Tanaka-Ueno, 2005, Mol. Phylogenet. Evol., 37: 733–742, found Humerana (exemplar Humerana miopus) to be imbedded phylogenetically within Hylarana sensu Frost et al. (2006) and therefore within Hylarana sensu Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1–13. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, confirmed the placement of Humerana miopus within Hylarana, although their adoption of a paraphyletic and antiquated taxonomy makes this difficult to appreciate. Hasan, Islam, Khan, Alam, Kurabayashi, Igawa, Kuramoto, and Sumida, 2012, Zool. Sci., Tokyo, 29: 162–172, also confirmed the placement of Humerana within Hylarana. Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 423, included one exemplar of Humerana (Humerana miopus) in their large molecular tree, which was recovered in a monophyletic group with Hylarana lateralis and Sylvirana guentheri. On the basis of a general lack of resolution of genera of the hylaranines in their tree, these authors placed Humerana into the synonymy of Hylarana. 

Chalcorana: Removed from the synonymy of Hylarana by Oliver, Prendini, Kraus, and Raxworthy, 2015, Mol. Phylogenet. Evol., 90: 187, where it had been placed by Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 3. See comment under Hylarana for access to relevant revisionary literature. Recovered as monophyletic, the sister taxon of Chalcorana, by Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 422. Regardless, the authors placed Chalcorana as a junior synonym of Hylarana due to the general lack of monophyletic genera within the hylaranine gr

Contained taxa (101 sp.):

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