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Megophryidae Bonaparte, 1850
Megalophreidina Bonaparte, 1850, Conspect. Syst. Herpetol. Amph.: 1 p. Type genus: Megalophrys Wagler, 1830 (= Megophrys Kuhl and Van Hasselt, 1822).
Megalophryinae — Fejérváry, 1922 "1921", Arch. Naturgesch., Abt. A,, 87: 25.
Megophryinae — Noble, 1931, Biol. Amph.: 492; Delorme, Dubois, Grosjean, and Ohler, 2006, Alytes, 24: 15.
Megalophryninae — Tamarunov, 1964, in Orlov (ed.), Osnovy Paleontologii, 12: 129.
Leptobrachiini Dubois, 1980, Bull. Mens. Soc. Linn. Lyon, 49: 471. Type genus: Leptobrachium Tschudi, 1838.
Megophryini — Dubois, 1980, Bull. Mens. Soc. Linn. Lyon, 49: 471; Delorme, Dubois, Grosjean, and Ohler, 2006, Alytes, 24: 15.
Leptobrachiinae — Dubois, 1983, Bull. Mens. Soc. Linn. Lyon, 52: 272; Dubois, 1983, Alytes, 2: 147; Mahony, Foley, Biju, and Teeling, 2017, Mol. Biol. Evol., 34: 752.
Oreolalaxinae Tian and Hu, 1985, Acta Herpetol. Sinica, Chengdu, N.S.,, 4 (3): 221. Type genus: Oreolalax Myers and Leviton, 1962. Considered a synonym of Leptobrachiinae by Dubois, 1987 "1986", Alytes, 5: 173–174; Fei and Ye, 2005, in Fei et al. (eds.), Illust. Key Chinese Amph.: 56.
Oreolalaginae — Dubois, 1987 "1986", Alytes, 5: 173–174. Justified emendation; Delorme, Dubois, Grosjean, and Ohler, 2006, Alytes, 24: 12.
Megophryidae — Ford and Cannatella, 1993, Herpetol. Monogr., 7: 94–117.
Xenophryini Delorme, Dubois, Grosjean, and Ohler, 2006, Alytes, 24: 17. Type genus Xenophrys Günther, 1864. Tribe.
Leptobrachiini — Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 131. Tribe.
Leptobrachiina — Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 131. Subtribe.
Leptolalagini — Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 131. Tribe.
Oreolalagina — Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 131. Subtribe.
Atympanophryini — Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 132. Type genus: Atympanophrys Tian and Hu, 1983. Tribe.
Brachytarsophryini — Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 132. Type genus: Brachytarsophrys Tian and Hu, 1983.
Grillitschiina Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 133. Type genus: Grillitschia Dubois, Ohler, and Pyron, 2021. Subtribe.
Ophryophryina Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 134. Type genus: Ophryophryne Boulenger, 1903. Subtribe.
Xenophryina — Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 134. Subtribe.
Common Names
Asian Toads (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 85; Li, Zhao, and Dong, 2010, Amph. Rept. Tibet: 14).
Litter Frogs (Iskandar, 1998, Amph. Java Bali: 35).
Leaf-litter Frogs (Grismer, 2012, Field Guide Amph. Rept. Seribuat Arch.: 39).
Asian Toadfrogs (Halliday and Adler, 2002, New Encyclop. Rept. Amph.: 82).
Short-legged Toads (Anders, 2002, in Schleich and Kästle (eds.), Amph. Rept. Nepal: 163).
Pseudomoustache Toads (Leptobrachiinae: Fei and Ye, 2016, Amph. China, 1: 449).
Distribution
Pakistan and western China east to the Philippines and the Greater Sunda Islands.
Comment
Rao and Yang, 1997, Asiat. Herpetol. Res., 7: 92–102, discussed karyology and phylogenetic relationships. Lathrop, 1997, Asiat. Herpetol. Res., 7: 68–79, and Haas, 2003, Cladistics, 19: 23-89, discussed relationships within Pelobatoidea and suggested that Megophryidae is the sister taxon of Pelobatidae and regarded them as subfamilies of Pelobatidae. Maglia, 1998, Sci. Pap. Nat. Hist. Mus. Univ. Kansas, 10: 1–19, suggested a relationship of (((Pelobatidae) (Pelodytidae)) Megophryidae). Dubois and Ohler, 1998, Dumerilia, 4: 1–32, discussed the classification of this taxon (ranked as a subfamily, Megophryinae, by Dubois and Ohler). Xie and Wang, 2000, Cultum Herpetol. Sinica, 8: 356–370, reviewed the taxonomy of this group (as part of Pelobatidae). Anders, 2002, in Schleich and Kästle (eds.), Amph. Rept. Nepal: 163–181, provided a key and accounts for the species in Nepal. Xu, 2005, Sichuan J. Zool., 24: 337–339, discussed karyotypes in Megophryidae and made taxonomic suggestions. Manthey and Grossmann, 1997, Amph. Rept. Südostasiens: 68–82, provided accounts and identification table for the species of the Sunda Shelf region. Dubois, 2005, Alytes, 23: 8, considered Megophryidae to be a subfamily of Pelobatidae, containing two tribes, Leptobrachiini and Megophryini. Li and Hu, 2005, Herpetol. Sinica, 10: 359–368, reported on karyological diversification within Megophryidae. 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, provided taxonomic history and a partial phylogenetic analysis and rejected subfamilies, although they did note that the clade (former Megophryinae) composed of Atympanophrys, Brachytarsophrys, Megophrys, Ophryophryne, and Xenophrys is presumably monophyletic. Delorme, Dubois, Grosjean, and Ohler, 2006, Alytes, 24: 6–21, provided a working taxonomy of the group based on a number of lines of previously published evidence, in which they formulated a system of generally paired taxa, one diagnosed on apomorphies, one on plesiomorphies. While this taxonomy did provide "chunks" to discuss these units, the plesiomorphic units have since been redelimited into monophyletic ones. The Delorme et al., 2006, system proposed three subfamilies: Leptobrachiinae (diagnosed on plesiomorphies shared with Megophryidae); Leptolalaginae (Leptobrachella, Leptolalax); and Megophryinae (containing Megophryini and Xenophryini [diagnosed on plesiomorphies shared with Megophryinae]). Within the aggregate Leptobrachiinae (= Megophryidae that are not members of Leptolalaginae and Megophryinae), two tribes were recognized, both suggested to be based on apomorphies: Leptobatrachiini (Leptobrachium) and Oreolalagini (Oreolalax and Scutiger). Within Megophryinae the tribe Megophryini (Borneophrys, Brachytarsophrys, and Megophrys) is suggested to rest on apomorphies, while Xenophryini (Ophryophryne + Xenophrys [which is also based on plesiomorphes, rendering it diagnostically equivalent to Megophryinae]) is based solely on characters primitive for Megophryinae. See other comments in the relevant generic records. Li, Guo, and Wang, 2011, Curr. Zool., Chengdu, 57: 93–100, provided a molecular analysis of Chinese species that suggested that Xenophrys is strongly paraphyletic with respect to at least Atympanophrys, Brachytarsophrys, and Ophryophryne (Megophrys sensu stricto not studied); this paraphyly has subsequently been corrected. Fu, Weadick, and Bi, 2007, J. Zool., London, 273: 315–325, reported on the molecular phylogenetics of Leptobrachium, Vibrissaphora, Oreolalax, and Scutiger, as did Rao and Wilkinson, 2008, Mol. Phylogenet. Evol., 46: 61–73, with denser taxon sampling and more data. Brown, Siler, Diesmos, and Alcala, 2010 "2009", Herpetol. Monogr., 23: 1–44, reported on molecular phylogenetics of Leptobrachium and additional provided evidence that the taxonomic arrangement of Delorme et al., 2006, is poorly correlated with phylogeny. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, confirmed the position of Megophryidae as the sister taxon of Pelobatidae. Blackburn and Wake, 2011, In Zhang (ed.), Zootaxa, 3148: 39–55, discussed briefly the taxonomic history of the group. Vitt and Caldwell, 2014, Herpetology, 4th Ed., provided a summary of life history, diagnosis, and taxonomy. Chen, Zhou, Poyarkov, Stuart, Brown, Lathrop, Wang, Yuan, Jiang, Hou, Chen, Suwannapoom, Nguyen, Duong, Papenfuss, Murphy, Zhang, and Che, 2017, Mol. Phylogenet. Evol., 106: 28–43, provided a detailed molecular study of phylogeny which resulted in the recognition of a monophyletic Atympanophrys, Megophrys, and Xenophrys. Mahony, Foley, Biju, and Teeling, 2017, Mol. Biol. Evol., 34: 744–771, subsequently provided a phylogenetic analysis focused on the megophryines, placed Atympanophrys and Xenophrys as subgenera of Megophrys, along with Ophryophryne, Brachytarsophrys, and Pelobatrachus (newly recognized as a taxon by them). These authors also provided estimated divergence times of taxa,and equated their enlarged Megophrys with Megophryinae and relegated other genera to Leptolabrachiinae. Chen, Zhou, Poyarkov, Stuart, Brown, Lathrop, Wang, Yuan, Jiang, Hou, Chen, Suwannapoom, Nguyen, Duong, Papenfuss, Murphy, Zhang, and Che, 2017, Mol. Phylogenet. Evol., 106: 28-43, reported on the molecular phylogenetics of the Megophryinae and partitioned Megophrys into five genera. Chen, Poyarkov, Suwannapoom, Lathrop, Wu, Zhou, Yuan, Jin, Chen, Liu, Nguyen, Nguyen, Duong, Eto, Nishikawa, Matsui, Orlov, Stuart, Brown, Rowley, Murphy, Wang, and Che, 2018, Mol. Phylogenet. Evol., 124: 162–171, reported on the molecular phylogenetics of Leptolalax and Leptobrachella, showing the latter is imbedded in the former and therefore placed Leptolalax into the synonymy of Leptobrachella. This was followed by Lyu, Zeng, Wang, Liu, Huang, Li, and Wang, 2021, Zootaxa, 4927: 9–40, who recognized 7 genera. Blotto, Pereyra, Grant, and Faivovich, 2020, Bull. Am. Mus. Nat. Hist., 443: 35, discussed possible morphological synapmorphies for the group, particularly of hand and foot morphology. Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 1–738, proposed a system of subfamilies, tribes, subtribes, and genera, dubiously suggested to increase communication. Elias-Costa, Araujo-Vieira, and Faivovich, 2021, Cladistics, 37: 498–517, discussed the evolution of submandibular musculature optimized on the tree of Jetz and Pyron, 2018, Nature Ecol. & Evol., 2: 850–858, which provided morphological synapomorphies of this taxon. Xiang, Zhou, Wei, Shu, Gu, and Jiang, 2024, Ecol. Evol., 4(e11687): 1–13, provided a molecular tree of Megophryidae, and a summary of base composition of mitogenomes for 26 species within the family.
Contained taxa (331 sp.):
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