Megophrys Kuhl and Van Hasselt, 1822

Class: Amphibia > Order: Anura > Family: Megophryidae > Subfamily: Megophryinae > Genus: Megophrys
5 species

Mogophrys Kuhl and Van Hasselt, 1822, Algemeene Konst-en Letter-Bode, 7: 102. Type species: Mogophrys montana Kuhl and Van Hasselt, 1822, by monotypy. See comment.

Megophrys Kuhl and Van Hasselt, 1822, Algemeene Konst-en Letter-Bode, 7: 104. Alternative spelling. See comment. 

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

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). 

Indonesian Horned Toads (Qi, Lyu, Wang, Mo, Zeng, Zeng, Dai, Li, Grismer, and Wang, 2021, Zootaxa, 5072: 430). 

Distribution

Java and Sumatra, Indonesia; records from Thailand southeast to Sumatra, Natuna, and Borneo apply to misidentifications. 

Comment

Given that at one time Megophrys was the generic name for all megophryines and and all currently recognized genera were at one time or another considered subgenera of Megophrys, the literature is very difficult to sort. Probably the best way to approach the literature is to start at the most recent and work backwards. See 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, all now redelimited and regarded as separate genera. Malkmus, Manthey, Vogel, Hoffmann, and Kosuch, 2002, Amph. Rept. Mount Kinabalu: 106, provided a key to Megophrys (now Megophrys, Pelobatrachus, and Sarawakiphrys) of Borneo. The intended original spelling of the generic name is unclear, Mogophrys on p. 102 with reference to the type species, Mogophrys montana, but Megophrys on p. 104 with reference to Megophrys monticola. Regardless, the current International Code of Zoological Nomenclature (1999), Art. 33.3.1, preserves the "prevailing usage", which is certainly "Megophrys", since the time of Kuhl and Van Hasselt, 1822, Isis von Oken, 10: 475, who would know what they intended in the earlier original publication. Li and Wang, 2008, Acta Zootaxon. Sinica, 33: 104–106, reviewed the species of China and concurred with the generic separation of Megophrys and XenophrysPyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, noted the paraphyly of Xenophrys (as then recognized) with respect to Megophrys, with such species as Xenophrys baluensis (now Pelobatrachus baluensis) as 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 into 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 (as then delimited), 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 their 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, 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 that time "Xenophrys" was a basal cloud within Megophryini and not monophyletic and the additional of Megophrys (which bears the oldest name) to that cloud hardly reduces the confusion and does not reduce the paraphyly at all. Wang, Zhang, Zhao, Sung, Yang, Pang, and Zhang, 2012, Zootaxa, 3546: 53–67, provided molecular evidence of the paraphyly of Xenophrys (as then delimited) 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 Xenophrys. Chen, Zhou, Poyarkov, Stuart, Brown, Lathrop, Wang, Yuan, Jiang, Hou, Chen, Suwannapoom, Nguyen, Duong, Papenfuss, Murphy, Zhang, and Che, 2017, Mol. Phylogenet. Evol., 106: 23–43, reported on a large-scale molecular analysis of the megophryines, which resulted in the re-recognition of a monophyletic Xenophrys and Atympanophrys, and the down-sizing of MegophrysAtympanophrys was subsequently 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, suggested that of their examplar species of 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 Ophryophryne. 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, 2017, Mol. Phylogenet. Evol., 106: 42, who provided a monophyletic taxon under this name. Brachytarsophrys was considered a subgenus of Megophrys by Dubois, 1987 "1986", Alytes, 5: 23, but retained as a genus by Zhao and Adler, 1993, Herpetol. China: 116. Rao and Yang, 1997, Asiat. Herpetol. Res., 7: 103–107, discussed systematics and distribution of the species of former BrachytarsophrysXie and Wang, 2000, Cultum Herpetol. Sinica, 8: 356–370, discussed evidence of the monophyly of Brachytarsophrys as well as character conflict regarding the monophyly of this taxon. Xu, 2005, Sichuan J. Zool., 24: 337–339, rejected Brachytarsophrys on karyological grounds. Fei, Hu, Ye, and Huang, 2009, Fauna Sinica, Amph. 2: 329–346, provided a key and accounts for the Chinese species. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, noted the paraphyly of Xenophrys (as then delimited) with respect to Brachytarsophrys, with such species as Xenophrys shapingensis and Xenophrys nankiangensis being more closely related to Brachytarsophrys than to other species of XenophrysFei 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. Kurniawan, Affandi, Wardani, Chomsy, Firdaus, and Kurnianto, 2017, J. Exp. Life Sci., 7: 76–83, reported on the molecular biogeography of several species, mostly unnamed, on Sumatra and Malaya. Mahony, Kamei, Teeling, and Biju, 2018, Zootaxa, 4523: 1–96, revised the Megophrys major complex in India, which involved the naming of four new species, the resurrection of Megophrys monticola, redelimitation of Megophrys parva, Megophrys glandulosa, and Megophrys major. One result is that much of the literature of Megophrys major may apply to other species, although this needs to be evaluated closely. Wang, Lyu, Liu, Liao, Zeng, Zhao, Li, and Wang, 2019, ZooKeys, 851: 113–164, added 6 new species of the subgenus Panophrys from montane southeastern China and provided a molecular tree of the species. Li, Zhang, Lyu, Wang, Li, Liu, Chen, Rao, Jin, Zhang, and Wang, 2020, Zool. Res., Kunming: 105–122, suggested that the nominal subgenera of Megophrys, recognized by Mahony, Foley, Biju, and Teeling, 2017, Mol. Biol. Evol., 34: 744–771, be recognized as genera. Mahony, Kamei, Teeling, and Biju, 2020, J. Nat. Hist., London, 54: 119–194, reported on specimens in northeastern India and Bangladesh that had previously had been identified as Megophrys parva, resulting in many reidentifications and the naming of three additional species. Lyu, Zeng, Wang, Liu, Huang, Li, and Wang, 2021, Zootaxa, 4927: 9–40, is the most recent summary of systematics and nomenclature. Munir, Nishikawa, Hamidy, and Smith, 2021, Zootaxa, 5057: 503–529, discussed the molecular and morphological systematics with special reference to Sumatra, Indonesia. Even though Megophrys montana, the type species of the genus, was found to be imbedded within Xenophrys in the tree of Portik, Streicher, and Wiens, 2023, Mol. Phylogenet. Evol., 188 (107907): 12, these authors did not make the taxonomic remedy. 

Contained taxa (5 sp.):

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