Megophrys Kuhl and Van Hasselt, 1822

Class: Amphibia > Order: Anura > Family: Megophryidae > Genus: Megophrys
76 species

Mogophrys Kuhl and Van Hasselt, 1822, Algemeene Konst-en Letter-Bode, 7: 104. Type species: Mogophrys montana Kuhl and Van Hasselt, 1822, by monotypy. See comment.

Megophrys — Kuhl and Van Hasselt, 1822, Isis von Oken, 10: 475. Incorrect subsequent spelling but protected as prevailing usage by Art. 33.3.1 of ICZN, 1999, Internatl. Code Zool. Nomencl., Ed. 4.

MegalophrysWagler, 1830, Nat. Syst. Amph.: 204. Incorrect subsequent spelling of Megophrys Kuhl and van Hasselt, 1822.

Ceratophryne Schlegel, 1858, Handl. Dierkd., 2: 56. Type species: Ceratophryne nasuta Schlegel, 1858, by subsequent designation of Gorham, 1966, Das Tierreich, 85: 15. See discussion by Dubois, 1980, Bull. Mens. Soc. Linn. Lyon, 49: 472. Synonymy by Inger, 1954, Fieldiana, Zool., 33: 223.

Xenophrys Günther, 1864, Rept. Brit. India: 414. Type species: Xenophrys monticola Günther, 1864 (= Leptobrachium parvum), by monotypy. 

Ophryophryne Boulenger, 1903, Ann. Mag. Nat. Hist., Ser. 7, 12: 186. Type species: Ophryophryne microstoma Boulenger, 1903, by monotypy.

Pelobatrachus Beddard, 1908 "1907", Proc. Zool. Soc. London, 1907: 909. Type species: Ceratophryne nasuta Schlegel, 1858, by monotypy.

Atympanophrys Tian and Hu, 1983, Acta Herpetol. Sinica, Chengdu, N.S.,, 2 (2): 43, 47. Type species: Megophrys shapingensis Liu, 1950, by original designation. Synonymy with Xenophrys by Delorme, Dubois, Grosjean, and Ohler, 2006, Alytes, 24: 18. Considered a subgenus of Megophrys by Dubois, 1987 "1986", Alytes, 5: 23.

Atympanophrys — Dubois, 1987 "1986", Alytes, 5: 23; Mahony, Foley, Biju, and Teeling, 2017, Mol. Biol. Evol., 34: 754. Treatment as a subgenus of Megophrys.

Panophrys Rao and Yang, 1997, Asiat. Herpetol. Res., 7: 98. Type species: Megophrys omeimontis, by original designation. Synonymy by Dubois and Ohler, 1998, Dumerilia, 4: 14.

Borneophrys Delorme, Dubois, Grosjean, and Ohler, 2006, Alytes, 24: 15. Type species: Megophrys edwardinae Inger, 1989, by original designation. Synonymy by Chen, Zhou, Poyarkov, Stuart, Brown, Lathrop, Wang, Yuan, Jiang, Hou, Chen, Suwannapoom, Nguyen, Duong, Papenfuss, Murphy, Zhang, and Che, 2016, Mol. Phylogenet. Evol., 106: 41. 

Xenophrys — Khonsue and Thirakhupt, 2001, Nat. Hist. J. Chulalongkorn Univ., 1: 75; Ohler, 2003, Alytes, 21: 23; Chen, Zhou, Poyarkov, Stuart, Brown, Lathrop, Wang, Yuan, Jiang, Hou, Chen, Suwannapoom, Nguyen, Duong, Papenfuss, Murphy, Zhang, and Che, 2016, Mol. Phylogenet. Evol., 106: 42. Treatment as a genus.

Liuophrys Fei, Ye, and Jiang in Fei and Ye, 2016, Amph. China, 1 : 614. Type species: Megophrys glandulosa Fei, Ye, and Huang, 1990. Provisionally considered a synonym of Megophrys due to appearing prior to the revision of Mahony, Foley, Biju, and Teeling, 2017, Mol. Biol. Evol., 34: 744–771.  

Borealophrys Fei, Ye, and Jiang in Fei and Ye, 2016, Amph. China, 1: 632. Type species: Megophrys nankiangensis Liu and Hu, 1966. Coined as a subgenus of Atympanophrys

Gigantophrys Fei, Ye, and Jiang in Fei and Ye, 2016, Amph. China, 1: 636. Type species: Megophrys giganticus Liu, Hu, and Yang, 1966. Coined as a subgenus of Atympanophrys

Boulenophrys Fei, Ye, and Jiang in Fei and Ye, 2016, Amph. China, 1: 641. Type species: Leptobrachium boettgeri Boulenger, 1899. 

Tianophrys Fei and Ye, 2016, Amph. China, 1: 676. Type species: Megophrys shuichengensis Tian, Gu, and Sun, 2000. Coined as a subgenus of Xenophrys

English Names

Asian Spadefoot Toads (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 86).

Asian Spadefoot Toads (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 87; Li, Zhao, and Dong, 2010, Amph. Rept. Tibet: 22).

Shaping Frogs (AtympanophrysFrank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 85).

Narrow-mouth Horned Toad (OphryophryneFei and Ye, 2016, Amph. China, 1: 729). 

Distribution

Tropical Asia from India and Bhutan to China and south to the Sundas and the Philippines. 

Comment

See comment under Megophryinae as well as discussion by Dubois and Ohler, 1998, Dumerilia, 4: 13–14. See comment under Megophrys montana. Dubois, 1987 "1986", Alytes, 5: 23, and Dubois and Ohler, 1998, Dumerilia, 4: 13–14, recognized four subgenera: Megophrys, Xenophrys, Brachytarsophrys and Atympanophrys. See comment under Ophryophryne. Malkmus, Manthey, Vogel, Hoffmann, and Kosuch, 2002, Amph. Rept. Mount Kinabalu: 106, provided a key to Megophrys (including Xenophrys) of Borneo. The original spelling of the generic name is clearly Mogophrys, not Megophrys. Nevertheless, the current Code of Zoological Nomenclature (1999), Art. 33.3.1, preserves the "prevailing usage", which is certainly "Megophrys". Li and Wang, 2008, Acta Zootaxon. Sinica, 33: 104–106, reviewed the species of China and concurred with the generic separation of Megophrys and Xenophrys. But, see comment under Megophryidae. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, in their study of Genbank sequences, noted the paraphyly of Xenophrys with respect to Megophrys, with such species as Xenophrys baluensis and being more closely related to Megophrys than to other species of Xenophrys. Rao and Yang, 1997, Asiat. Herpetol. Res., 7: 98, placed a number of species in Panophrys, but did not address all species in Megophrys, rendering the content and distribution of both nominal genera uncertain. Jiang, Yuan, Xie, and Zheng, 2003, Zool. Res., Kunming, 24: 246, suggested on the basis of DNA sequence data and karyotype that Atympanophrys and Brachytarsophrys are imbedded within Xenophrys, more closely related to such species as Megophrys minorMegophrys brachykolos, and Megophrys omeimontis, than Megophrys glanulosaDelorme, Dubois, Grosjean, and Ohler, 2006, Alytes, 24: 6–21, suggested on the basis of morphological comparisons that Brachytarsophrys is not part of this phylogenetic assemblage and that Xenophrys is not diagnosable against the backdrop of Megophryini. Fei, Ye, Huang, Jiang, and Xie, 2005, in Fei et al. (eds.), Illust. Key Chinese Amph.: 77, regarded Atympanophrys as the Megophrys shapingensis group (in which they also included Xenophrys nankiangensis) and provided a key to Chinese species under this nomenclature. Xu, 2005, Sichuan J. Zool., 24: 337–339, regarded Atympanophrys as valid on karyological grounds. Li and Wang, 2008, Acta Zootaxon. Sinica, 33: 104–106, reviewed the species of China and concurred with the generic separation of Megophrys and XenophrysFei, Hu, Ye, and Huang, 2009, Fauna Sinica, Amph. 2: 346–471, provided a key and accounts for the species of China (as Megophrys). Mahony, 2011, Zootaxa, 2734: 23–39, and more formally by Mahony, Teeling, and Biju, 2013, Zootaxa, 3722: 144, suggested that Xenophrys should be considered a junior synonym of Megophrys. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543-583, noted the paraphyly of Xenophrys with respect to Megophrys and Brachytarsophrys (and did not address Borneophrys), with such species as Megophrys shapingensis and Megophrys nankiangensis being more closely related to Brachytarsophrys than to other species of Xenophrys and Xenophrys baluensis being more closely related to Megophrys. So, at present "Xenophrys" is a basal cloud within Megophryini and not monophyletic and the additional of Megophrys (which bears the older name) to that cloud hardly reduces the confusion and does not reduce the paraphyly at all, but this arrangement is followed here because that appears to be the predominant usage of those currently working on the group. Wang, Zhang, Zhao, Sung, Yang, Pang, and Zhang, 2012, Zootaxa, 3546: 53-67, provided molecular evidence of the paraphyly of Xenophrys with respect to Ophryophryne, but did not make the nomenclatural remedy. Wang, Zhao, Yang, Zhou, Chen, and Liu, 2014, PLoS One, 9(4)(e93075): 1–15, reported on the molecular phylogenetics, morphology, and bioacoustics of a group of nominal Xenophrys in southern China. Orlov, Poyarkov, and Nguyen, 2015, Russ. J. Herpetol., 22: 206–218, provided an identification key and extensive comments on characteristics of the species of Vietnam and explicitly addressed the apparent paraphyly of the genus. Chen, Zhou, Poyarkov, Stuart, Brown, Lathrop, Wang, Yuan, Jiang, Hou, Chen, Suwannapoom, Nguyen, Duong, Papenfuss, Murphy, Zhang, and Che, 2016, Mol. Phylogenet. Evol., 106: 23–43, reported on a large-scale molecular analysis of the megophryines, which resulted in the re-recognition of Xenophrys and Atympanophrys, and the down-sizing of Megophrys.Atympanophrys was resurrected as a genus in the revision by Chen, Zhou, Poyarkov, Stuart, Brown, Lathrop, Wang, Yuan, Jiang, Hou, Chen, Suwannapoom, Nguyen, Duong, Papenfuss, Murphy, Zhang, and Che, 2016, Mol. Phylogenet. Evol., 106: 28–43, but sequently treated as a subgenus of Megophrys by Mahony, Foley, Biju, and Teeling, 2017, Mol. Biol. Evol., 34: 754. Ophryophryne was considered by Dubois, 1980, Bull. Mens. Soc. Linn. Lyon, 49: 473, to be ranked at the level of subgenus (of Megophrys), but ranked as a genus subsequently by Dubois, 1987 "1986", Alytes, 5: 23. The phylogenetic position of this taxon was discussed by Rao and Yang, 1997, Asiat. Herpetol. Res., 7: 92-102. Ohler, 2003, Alytes, 21: 23-44, revised the genus and provided a key to the species. Fei, Ye, Huang, Jiang, and Xie, 2005, in Fei et al. (eds.), Illust. Key Chinese Amph.: 87, provided a key to the Chinese species of OphryophryneDelorme, Dubois, Grosjean, and Ohler, 2006, Alytes, 24: 17, provided evidence that Ophryophryne is a monophyletic derivative of a paraphyletic XenophrysFei, Hu, Ye, and Huang, 2009, Fauna Sinica, Amph. 2: 472-481, provided a key and accounts for the Chinese species. See comment under Megophryidae. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543-583, in their study of Genbank sequences, suggested that of their examplar species Ophryophryne is the sister taxon of a monophyletic group composed of "Xenophrys", Megophrys, and BrachytarsophrysWang, Zhang, Zhao, Sung, Yang, Pang, and Zhang, 2012, Zootaxa, 3546: 53-67, provided molecular evidence of the paraphyly of Xenophrys with respect to Ophryophryne, but did not make the nomenclatural remedy. Ophryophryne was treated as a subgenus of Megophrys by Mahony, Foley, Biju, and Teeling, 2017, Mol. Biol. Evol., 34: 744–771. Poyarkov, Duong, Orlov, Gogoleva, Vassilieva, Nguyen, Nguyen, Nguyen, Che, and Mahony, 2017, ZooKeys, 672: 49–120, reported on phylogenetics and morphological and acoustic diversity of OphryophryneXenophrys was placed in the synonymy with Leptobrachium by Boulenger, 1889, Ann. Mus. Civ. Stor. Nat. Genova, Ser. 2, 7: 748; with Megalophrys byBoulenger, 1908, Proc. Zool. Soc. London, 1908: 407. Considered a subgenus of Megophrys by Dubois and Ohler, 1998, Dumerilia, 4: 14, and a synonym, of Megophrys by Mahony, Teeling, and Biju, 2013, Zootaxa, 3722: 144. Resurrected from synonymy by Chen, Zhou, Poyarkov, Stuart, Brown, Lathrop, Wang, Yuan, Jiang, Hou, Chen, Suwannapoom, Nguyen, Duong, Papenfuss, Murphy, Zhang, and Che, 2016, Mol. Phylogenet. Evol., 106: 42, who provided a monophyletic taxon under this name. Fei and Ye, 2016, Amph. China, 1: 611–736, reviewed the species of China under a variety of generic and subgeneric names, providing accounts, photographs, and dot maps; because this volume was formulated prior to the revision of Mahony, Foley, Biju, and Teeling, 2017, Mol. Biol. Evol., 34: 744–771, I follow the latter taxonomy although I do expect that Megophrys as currently recognized will be generically partitioned. 

Contained taxa (76 sp.):

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