Boulenophrys Fei, Ye, and Jiang, 2016

Class: Amphibia > Order: Anura > Family: Megophryidae > Subfamily: Megophryinae > Genus: Boulenophrys
67 species

Panophrys Rao and Yang, 1997, Asiat. Herpetol. Res., 7: 98. Type species: Megophrys omeimontis, by original designation. Preoccupied by Panophrys Dujardin, 1840, as first noted by Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 524. 

Boulenophrys Fei, Ye, and Jiang in Fei and Ye, 2016, Amph. China, 1: 641. Type species: Leptobrachium boettgeri Boulenger, 1899. Treated as senior to Tianophrys Fei and Yei, 2016, by Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 469, 524, and 679, as first revisors. 

Tianophrys Fei and Ye, 2016, Amph. China, 1: 676. Type species: Megophrys shuichengensis Tian, Gu, and Sun, 2000. Coined as a subgenus of Xenophrys. Synonymy with Panophrys (by implication) of Lyu, Zeng, Wang, Liu, Huang, Li, and Wang, 2021, Zootaxa, 4927: 9–40.

PanophrysMahony, Foley, Biju, and Teeling, 2017, Mol. Biol. Evol., 34: 755. Treatment as a subgenus of Megophrys

Panophrys — Lyu, Zeng, Wang, Liu, Huang, Li, and Wang, 2021, Zootaxa, 4927: 14. Treatment as a genus. 

Boulenophrys — Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 469, 524, and 679. 

English Names

Chinese Horned Toad (Qi, Lyu, Wang, Mo, Zeng, Zeng, Dai, Li, Grismer, and Wang, 2021, Zootaxa, 5072: 431). 

Lesser Horned Toad (Dinesh, Radhakrishnan, Deepak, and Kulkarni, 2023, Fauna India Checklist, vers. 5.0 : 6).

Distribution

Most of China except for Tibet, extending south into Vietnam, Laos, Thailand, Myanmar, with less certainty to Bhutan and northeastern India. 

Comment

Removed from the synonymy of Megophrys by Lyu, Zeng, Wang, Liu, Huang, Li, and Wang, 2021, Zootaxa, 4927: 14, where it had been placed variously as a subgenus of Megophrys by Dubois and Ohler, 1998, Dumerilia, 4: 14, Mahony, Foley, Biju, and Teeling, 2017, Mol. Biol. Evol., 34: 744–771, or as distinct genus 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: 28–43. Wu, Suwannapoom, Poyarkov, Chen, Pawangkhanant, Xu, Jin, Murphy, and Che, 2019, Zool. Res., Kunming, 40: 564–574, discussed comparative characteristics within the Panophrys minor complex and provided a molecular tree of a segment of the diversity (then in Xenophrys). Luo, Wang, Wang, Lu, Wang, Deng, and Zhou, 2021, ZooKeys, 1047: 27–60, provided a ML tree based on two mtDNA loci for species within the genus as well as representatives of other genera within Megophryidae. Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 469, 524, and 679, first recognized that Panophrys is proccupied and employed Boulenophrys as the generic name rather than Tianophrys under the Principle of First Revisor. This was discussed in detail by Qi, Lyu, Wang, Mo, Zeng, Zeng, Dai, Li, Grismer, and Wang, 2021, Zootaxa, 5072: 401–438, who provided a checklist of the entire genus and otherwise review the recent taxonomic literature of Megophryinae. These authors provided a system of three species groups and one collection of incertae sedis species summarized here: (1) Boulenophrys boettgeri group: Boulenophrys baishanzuensis, Boulenophrys baolongensis, Boulenophrys brachykolos, Boulenophrys caudoprocta, Boulenophrys boettgeri, Boulenophrys acuta, Boulenophrys cheni, Boulenophrys congjiangensis, Boulenophrys daiyunensis, Boulenophrys daoji, Boulenophrys dongguanensis, Boulenophrys huangshanensis, Boulenophrys insularis, Boulenophrys jinggangensis, Boulenophrys jiulianensis, Boulenophrys kuatunensis, Boulenophrys leishanensis, Boulenophrys liboensis, Boulenophrys lini, Boulenophrys lishuiensis, Boulenophrys mirabilis, Boulenophrys mufumontana, Boulenophrys nankunensis, Boulenophrys nanlingensis, Boulenophrys obesa, Boulenophrys ombrophila, Boulenophrys sanmingensis, Boulenophrys shimentaina, Boulenophrys shunhuangensis, Boulenophrys tongboensis, Boulenophrys tuberogranulata, Boulenophrys wugongensis, Boulenophrys wushanensis, Boulenophrys xiangnanensis, Boulenophrys xianjuensis, Boulenophrys yangmingensis, Boulenophrys yaoshanensis, Boulenophrys yingdeensis, and Boulenophrys yunkaiensis; (2) Boulenophrys minor group: Boulenophrys jiangi, Boulenophrys minor, and Boulenophrys chishuiensis; (3) Boulenophrys omeimontis group: Boulenophrys angka, Boulenophrys anlongensis, Boulenophrys binchuanensis, Boulenophrys binlingensis, Boulenophrys caobangensis, Boulenophrys daweimontis, Boulenophrys jingdongensis, Boulenophrys lushuiensis, and Boulenophrys omeimontis; and incertae sedis within Boulenophrys: Boulenophrys fansipanensis, Boulenophrys frigida, Boulenophrys hoanglienensis, and Boulenophrys shuichengensis. Dubois, 2021, Bionomina, 24: 38–50, criticized Poyarkov, Nguyen, Popov, Geissler, Pawangkhanant, Neang, Suwannapoom, and Orlov, 2021, Russ. J. Herpetol., 28 (3A): 36 (June), for employing the name Panophrys rather than Boulenophrys, first shown to be the appropriate generic name by Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 524. The tone of Dubois' attack was hardly appropriate given that Poyarkov's paper may well have already been in press by the time Dubois et al.'s paper appeared (DRF). Wang, Zeng, Lyu, Qi, Liu, Chen, Lu, Xiao, Lin, Chen, and Wang, 2022, Zootaxa, 5099: 91–119, provided a general discussion of the systematics within the genus, including an mtDNA tree. Lyu, Qi, Wang, Zhang, Zhao, Zeng, Wan, Yang, Mo, and Wang, 2023, Zool. Res., Kunming, 44: 380–450, provided an identification key and accounts for the Chinese species. 

Contained taxa (67 sp.):

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