Dicroglossidae Anderson, 1871
Dicroglossidae Anderson, 1871, J. Asiat. Soc. Bengal, 40: 38. Type genus: see comment.
Occydozyginae Fei, Ye, and Huang, 1990, Key to Chinese Amph.: 123. Type genus: Occidozyga Kuhl and Van Hasselt, 1822.
Paini Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 317. Type genus: Paa Dubois, 1975. Synonymy by Roelants, Jiang, and Bossuyt, 2004, Mol. Phylogenet. Evol., 31: 734-735.
Limnonectini Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 315. Type genus: Limnonectes Fitzinger, 1843.
Occydozyginae — Dubois, Ohler, and Biju, 2001, Alytes, 19: 55.
Occydozygini — Dubois, 2005, Alytes, 23: 16.
Annandini — Fei, Ye, and Jiang, 2010, Herpetol. Sinica, 12: 19. Type genus: Annandia Dubois, 1992. See comment.
Hoplobatrachini Fei, Ye, and Jiang, 2010, Herpetol. Sinica, 12: 19. Type genus: Hoplobatrachus Peters, 1863. See comment.
Ingeranini Fei, Ye, and Jiang, 2010, Herpetol. Sinica, 12: 19. Type genus: Ingerana Dubois, 1986. See comment.
Nannophryini Fei, Ye, and Jiang, 2010, Herpetol. Sinica, 12: 19. Type genus: Nannophrys Günther, 1869. See comment.
Quasipaini Fei, Ye, and Jiang, 2010, Herpetol. Sinica, 12: 22. Type genus: Quasipaa Dubois, 1992. See comment.
Fork-tongued frogs (O'Shea, Sanchez, Heacox, Kathriner, Carvalho, Ribeiro, Soares, Araujo, and Kaiser, 2012, Asian Herpetol. Res., Ser. 2, 3: 114-126).
Northwestern and subsaharan Africa, southern Arabian Peninsula, Pakistan and India to Afghanistan, Nepal, Malaya, and Sri Lanka; east through Nepal and Myanmar to western and southern China, Indochina, and islands of the Sunda Shelf; Philippines; Japan; reported in Papua New Guinea.
Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 317, diagnosed Dicroglossinae in the sense of including Conrauinae and excluding Paini. Marmayou, Dubois, Ohler, Pasquet, and Tillier, 2000, C. R. Acad. Sci., Ser. 3, Paris, 323: 287–297, and Delorme, Dubois, Kosuch, and Vences, 2004, Alytes, 22: 53–Occidozyga and Phrynoglossus in the subfamily Occidozyginae. Grosjean, Vences, and Dubois, 2004, Biol. J. Linn. Soc., 81: 171–181, suggested on the basis of several mt and nuDNA loci that Euphlyctis (Dicroglossini) is the sister taxon of Hoplobatrachus (Limnonectini) with Fejervarya, Sphaerotheca, Nannophrys, and Limnonectes forming more distant relations, a result consistent with the tree of Roelants, Jiang, and Bossuyt, 2004, Mol. Phylogenet. Evol., 31: 735. Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 315, also recognized a tribe Limnonectini diagnosed nearly identically with Conrauini (now in Petropedetidae), differing only in the larval keratodont formula. Nominal genera contained in this group occur from tropical Africa with most taxonomic diversity in tropical Asia: Hoplobatrachus, Limnonectes, and Fejervarya. In addition, Marmayou, Dubois, Ohler, Pasquet, and Tillier, 2000, C. R. Acad. Sci., Ser. 3, Paris, 323: 287–297, and Delorme, Dubois, Kosuch, and Vences, 2004, Alytes, 22: 53–64, suggested on the basis of mtDNA evidence that Sphaerotheca (formerly in Tomopterninae—Dubois, 1987 "1986", Alytes, 5: 56–57) and Taylorana (now a synonym of Limnonectes; originally considered to be a member of Limnonectini [Dubois, 1987 "1986", Alytes, 5: 63] but subsequently transferred to Ceratobatrachini by Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 314) are in Limnonectini. Roelants, Jiang, and Bossuyt, 2004, Mol. Phylogenet. Evol., 31: 735, also placed Nannophrys in Dicroglossinae ; by implication on the basis of mt and nuDNA evidence; it was previously assigned to Ranixalini by Dubois, 1987 "1986", Alytes, 5: 66, and Dicroglossini by Dubois, Ohler, and Biju, 2001, Alytes, 19: 56. Dubois, Ohler, and Biju, 2001, Alytes, 19: 55, implied on the basis of various published and unpublished mtDNA data that Euphlyctis (formerly in his Dicroglossini), Fejervarya, Hoplobatrachus, Minervarya, Nannophrys, and Sphaerotheca (formerly in his Limnonectini) should be included in a reconstituted Dicroglossini. Dubois, 2005, Alytes, 23: 16, recognized four tribes, Dicroglossini (for Euphlyctis, Fejervarya, Minervarya, Nannophrys, and Sphaerotheca), Limnonectini (for Limnonectes, as well as some taxa considered by most authors to be synonyms of Limnonectes), Occidozygini (for Occidozyga and Phrynoglossus), and Paini (for Chaparana, Nanorana, and Quasipaa). 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, discussed the taxonomy of this group as well as its convoluted taxonomic history and recognized two subfamilies, Dicroglossinae and Occidozyginae, rejected Paini as paraphyletic and provided a partial revision. Ohler and Dubois, 2006, Zoosystema, 28: 769–784, provided a phylogenetic analysis of the former Paini and suggested that it is monophyletic, as long as Annandia delacouri is excluded from the group, and assuming that one species of Hoplobatrachus, one species of Euphlyctis, one species of Limnonectes, and one species of Fejervarya as outgroups provides an appropriate test for this proposition. Kurabayashi, Kuramoto, Joshy, and Sumida, 2005, Zool. Sci., Tokyo, 22: 525–534, reported on a moleclar phylogeny of Indian species of Fejervarya, Hoplobatrachus, and Euphlyctis. See comment under Ingerana (Ceratobatrachidae) of which the type species, Ingerana tenasserimensis belongs in Dicroglossidae, although it is not clear what component of "Ingerana" should remain in Ceratobatrachidae, beyond "Ingerana" baluensis. Vitt and Caldwell, 2009, Herpetology, 3rd Ed.: 475, provided a general taxonomic account and map as part of a much more general and extensive overview of amphibian biology. Wiens, Sukumaran, Pyron, and Brown, 2009, Evolution, 63: 1217–1231, suggested considerable phylogenetic structure within Dicroglossidae (their Dicroglossinae), although the phylogenetic placement of this taxon as the sister taxon of Micrixalinae lacks any confidence measure suggesting that this placement is poorly supported. Bossuyt and Roelants, 2009, in Hedges and Kumar (eds.), Timetree of Life: 357–364, considered this taxon a distinct family based on its Mesozoic origin. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, in their study of Genbank sequences confirmed the monophyly of this taxon and its subfamilies and provided an estimate of the phylogeny of a number of exemplar species as well as providing a non-monophyletic taxonomy. See comments in the subsidiary accounts for these results. Blackburn and Wake, 2011, In Zhang (ed.), Zootaxa, 3148: 39–55, discussed briefly the taxonomic history of the group. Chen, Wang, Liu, Xie, and Jiang, 2011, Curr. Zool., Chengdu, 57: 785–805, on the basis of 11 protein-coding mtDNA genes, suggested that Dicroglossidae is polyphyletic, the two subfamilies not having a lot to do with each other, atlhoguh the confidence measures were low. Fei, Ye, and Jiang, 2010, Herpetol. Sinica, 12: 1–43, provided a new classification of Dicroglossidae (their Dicroglossidae and Occidozygidae), suggested to be phylogenetic but apparently more of an attempt to rehabilitate a gradistic classification that is here discussed, but not implemented in the catalogue for reason of its explicitly nonmonophyletic groups—at least when cast upon the phylogenetic trees of Che, Hu, Zhou, Murphy, Papenfuss, Chen, Rao, Li, and Zhang, 2009, Mol. Phylogenet. Evol., 50: 59–73, and Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, and its lack of new data to address previously documented paraphyly. Within their Dicroglossidae these authors recognized three subfamilies: Dicroglossinae, Painae, and Limnonectinae, which phylogenetically are in the topology Dicroglossinae + (Painae + Limnonectinae). Within Dicroglossinae they recognized a nonsymmetrical set of monophyletic tribes, Nannophryini (for Nannophrys), Dicroglossinae (for Euphlyctis), Hoplobatrachini (for Hoplobatrachus), and Fejervaryini (for Sphaerotheca, Fejervarya [including what is now Zakerana], and Minervarya [see comment under that taxon record]). Within Limnonectinae they recognized two tribes: Limnonectini (to include Limnonectes and Elachyglossa [although recognition of this taxon requires the paraphyly of Limnonectes]) and Annandiini (for a resurrected Annandia, although this taxon was placed in the synonymy of Quasipaa Che, Hu, Zhou, Murphy, Papenfuss, Chen, Rao, Li, and Zhang, 2009, Mol. Phylogenet. Evol., 50: 69]). Within Paini, the authors recognized two tribes. The first, Nanoranini, was made to contain Ombrana, Allopaa, and Chrysopaa (none of which has even been submitted to any kind of phylogenetic analysis and have only retained by other authors solely as a statement of ignorance), and a number of resurrected and new genera: Maculopaa (likely monophyletic), Nanorana (redlimited and likely monophyletic), Paa (of uncertain monophyly), Feirana (likely paraphyletic with respect to Unculurana and Chaparana), Unculurana (monotypic), Chaparana (likely monophyletic). Within Quasipaini they placed Quasipaa(rendered paraphyletic by the recognition of Yerana) and Yerana (which had been placed in the synonymy of Quasipaa by implication of Che, Hu, Zhou, Murphy, Papenfuss, Chen, Rao, Li, and Zhang, 2009, Mol. Phylogenet. Evol., 50: 66). Within their Occidozygidae, Fei, Ye, and Jiang, 2010, Herpetol. Sinica, 12: 1–43, recognized two subfamilies: Occidozyginae and Liuraninae. Within Occidozyginae, the authors recognized Occidozyga and Phrynoglossus (although recognition of Phrynoglossus in their sense likely renders Occidozyga paraphyletic [note also that Phrynoglossus, and therefore Occidozyga, is paraphyletic with respect to Ingerana on the tree of Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583]. See comment under Ingerana borealis). Hu, Xie, Li, and Jiang, 2011, PLoS One, 6(5): e19817: 1–10, reported on elevational patterns of species richness, range, and body size of Allopaa, Chrysopaa, and Quasipaa. Within Ingeranini (likely rendering Occidozyginae paraphyletic, see above) the authors recognized Ingerana (restricted to Ingerana charlesdarwini, Ingerana tasanae, and Ingerana tenasserimensis) and Taylorana (although recognition of this genus renders Limnonectes (!) paraphyletic according to Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583. They also recognize a tribe Liuranini for Liurana, resurrected from the synonymy of Ingerana, although this cluster of species has yet to be subjected to any kind of phylogenetic analysis. As with the taxonomy provided by these authors in the same paper for Ranidae, I provisionally set it aside, until much of the paraphyly provided by these authors can be addressed by others. Vitt and Caldwell, 2013, Herpetology, 4th Ed., provided a summary of life history, diagnosis, and taxonomy. Ohler and Dubois, 2014, Zootaxa, 3838: 590–594, found that the name Dicroglossidae Anderson, 1871, is based on a misspelling of Discoglossus Gunther, 1864. Rather than apply the next oldest name, which would by Occydozygidae Fei, Ye, and Huang, 1990, Key to Chinese Amph.: 123, Ohler and Dubois suggested to retain the name in general usage until they could appeal to the ICZN for a ruling to preserve the name. I (DRF) have followed that advice. Li, Zhang, Wu, Xue, Yan, and Wu, 2014, Amphibia-Reptilia, 35: 331-343, reported on the molecular phylogenetics of the group on the basis of 12 examplars and 13 mtDNA protein-coding genes.
Contained taxa (186 sp.):
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