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III. Genus Atympanophrys Tian & Hu, 1983 Suggested common name: Hidden-tympanum horned toads (in English) / ĿHDzfl (in Chinese) Four species: Atympanophrys shapingensis (Liu, 1950) (type species); Atympanophrys gigantica (Liu, Hu & Yang, 1960); Atympanophrys nankiangensis (Liu & Hu, 1966); Atympanophrys wawuensis (Fei, Jiang & Zheng, 2001)., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

2) Boulenophrys minor group Three species: Boulenophrys minor (Stejneger, 1926); Boulenophrys chishuiensis (Xu, Li, Liu, Wei & Wang, 2020) comb. nov.; Boulenophrys jiangi (Liu, Li, Wei, Xu, Cheng, Wang & Wu, 2020) comb. nov.., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 431, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

3) Boulenophrys omeimontis group Fifteen species: Boulenophrys omeimontis (Liu, 1950); Boulenophrys anlongensis (Li, Lu, Liu and Wang, 2020) comb. nov.; Boulenophrys angka (Wu, Suwannapoom, Poyarkov, Pawangkhanant, Xu, Jin, Murphy & Che, 2019) comb. nov.; Boulenophrys binchuanensis (Ye & Fei, 1995); Boulenophrys binlingensis (Jiang, Fei & Ye, 2009) comb. nov.; Boulenophrys caobangensis (Nguyen, Pham, Nguyen, Luong & Ziegler, 2020) comb. nov.; Boulenophrys daweimontis (Rao & Yang, 1997) comb. nov.; Boulenophrys jingdongensis (Fei & Ye, 1983) comb. nov.; Boulenophrys lushuiensis (Shi, Li, Zhu, Jiang, Jiang & Wang, 2021) comb. nov.; Boulenophrys palpebralespinosa (Bourret, 1937) comb. nov.; Boulenophrys qianbeinsis (Su, Shi, Wu, Li, Yao, Wang & Li, 2020) comb. nov.; Boulenophrys rubrimera (Tapley, Cutajar, Mahony, Chung, Dau, Nguyen, Luong & Rowley, 2017) comb. nov.; Boulenophrys sangzhiensis (Jiang, Ye & Fei, 2008) comb. nov.; Boulenophrys spinata (Liu & Hu, 1973); Boulenophrys wuliangshanensis (Ye & Fei, 1995)., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 431, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

1) Boulenophrys boettgeri group Thirty-nine species: Boulenophrys boettgeri (Boulenger, 1899) (type species); Boulenophrys acuta (Wang, Li & Jin, 2014); Boulenophrys baishanzuensis (Wu, Li, Liu, Wang & Wu, 2020) comb. nov.; Boulenophrys baolongensis (Ye, Fei & Xie, 2007); Boulenophrys brachykolos (Inger & Romer, 1961); Boulenophrys caudoprocta (Shen, 1994) comb. nov.; Boulenophrys cheni (Wang & Liu, 2014); Boulenophrys congjiangensis (Luo, Wang, Wang, Lu, Wang, Deng & Zhou, 2021) comb. nov.; Boulenophrys daiyunensis (Lyu, Wang & Wang, 2021) comb. nov.; Boulenophrys daoji (Lyu, Zeng, Wang & Wang, 2021) comb. nov.; Boulenophrys dongguanensis (Wang & Wang, 2019) comb. nov.; Boulenophrys huangshanensis (Fei & Ye, 2005); Boulenophrys insularis (Wang, Liu, Lyu, Zeng & Wang, 2017) comb. nov.; Boulenophrys jinggangensis (Wang, 2012); Boulenophrys jiulianensis (Wang, Zeng, Lyu & Wang, 2019) comb. nov.; Boulenophrys kuatunensis (Pope, 1929); Boulenophrys leishanensis (Li, Xu, Liu, Jiang, Wei & Wang, 2018) comb. nov.; Boulenophrys liboensis (Zhang, Li, Xiao, Li, Pan, Wang, Zhang & Zhou, 2017) comb. nov.; Boulenophrys lini (Wang & Yang, 2014); Boulenophrys lishuiensis (Wang, Liu & Jiang, 2017); Boulenophrys mirabilis (Lyu, Wang & Zhao, 2020) comb. nov.; Boulenophrys mufumontana (Wang, Lyu & Wang, 2019) comb. nov.; Boulenophrys nankunensis (Wang, Zeng & Wang, 2019) comb. nov.; Boulenophrys nanlingensis (Lyu, Wang, Liu & Wang, 2019) comb. nov.; Boulenophrys obesa (Wang, Li & Zhao, 2014); Boulenophrys ombrophila (Messenger & Dahn, 2019) comb. nov.; Boulenophrys sanmingensis (Lyu & Wang, 2021) comb. nov.; Boulenophrys shimentaina (Lyu, Liu, & Wang, 2020) comb. nov.; Boulenophrys shunhuangensis (Wang, Deng, Liu, Wu & Liu, 2019) comb. nov.; Boulenophrys tongboensis (Wang & Lyu, 2021) comb. nov.; Boulenophrys tuberogranulatus (Shen, Mo & Li, 2010); Boulenophrys wugongensis (Wang, Lyu & Wang, 2019) comb. nov.; Boulenophrys wushanensis (Ye & Fei, 1995); Boulenophrys xiangnanensis (Lyu, Zeng & Wang, 2020) comb. nov.; Boulenophrys xianjuensis (Wang, Wu, Peng, Shi, Lu & Wu, 2020) comb. nov.; Boulenophrys yangmingensis (Lyu, Zeng, & Wang, 2020) comb. nov.; Boulenophrys yaoshanensis Qi, Mo, Lyu, Wang & Wang sp. nov.; Boulenophrys yingdeensis Qi, Lyu, Wang & Wang sp. nov.; Boulenophrys yunkaiensis Qi, Wang, Lyu & Wang sp. nov.., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 431, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

1) Brachytarsophrys carinense group Two species: Brachytarsophrys carinensis (Boulenger, 1889) (type species); Brachytarsophrys intermedia (Smith, 1921)., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

Boulenophrys yunkaiensis sp. nov. Qi, Wang, Lyu & Wang Yunkai Horned Toad / yun kai jiao chan (云开fflDz) Figures 6, 7C Chresonymy. Megophrys sp 32 (SYS a004637–4638, 4694)— Liu et al. 2018 Holotype. SYS a004637 (Figs. 6A, 7C), adult male, collected on 14 April 2016 by Jian Wang, Zhi-Tong Lyu and Ying-Yong Wang from the Yunkaishan Nature Reserve (22.2758°N, 111.1952°E; ca 950 m a.s.l.), Xinyi City, Guangdong Province, China. Paratypes. Eight adult specimens from the same locality as the holotype: males SYS a004636 and SYS a004638 collected on 14 April 2016 by Jian Wang, Zhi-Tong Lyu and Ying-Yong Wang; male SYS a004662/ CIB 116085 and females SYS a004659 (Figs. 6B) and SYS a004660 collected on 15 April 2016 by Jian Wang, Zhi-Tong Lyu and Ying-Yong Wang; female SYS a004691 collected on 16 April 2016 by Jian Wang, Zhi-Tong Lyu and Ying-Yong Wang; males SYS a004986 and SYS a004987 collected on 28 June 2016 by Jian Wang. Etymology: The specific epithet yunkaiensis refers to its type locality, the Yunkaishan Nature Reserve, western Guangdong, China. Diagnosis. (1) Small body size, SVL 35.3–40.0 mm (37.6 ± 1.7, N = 6) in adult males and SVL 45.3–46.1 mm (45.8 ± 0.4, N = 3) in adult females; (2) snout rounded in dorsal view; (3) tympanum boundary clear, ED/TD 1.68–1.91 in males, 1.47–1.80 in females; (4) presence of vomerine ridge and absence of vomerine teeth; (5) margin of tongue rounded, not notched behind; (6) hindlimbs slender, heels overlapping or just meeting and tibio-tarsal articulation reaching forward between tympanum to posterior corner of eye when leg stretched forward; (7) tibia 0.40–0.48 of SVL and foot 0.60–0.68 of SVL in males, while tibia 0.42–0.46 of SVL and foot 0.59–0.66 of SVL in female; (8) a subarticular tubercle present at the base of each fingers; (9) toes without lateral fringes and with only rudimentary webbing; (10) presence of small horn-like tubercle at the edge of upper eyelid; (11) surface around cloaca with large tubercles bearing tiny spines; (12) dorsal skin rough with small granules, a discontinuous “X” or “Y”-shaped ridge with two discontinuous dorsolateral ridges on two side of dorsum; (13) sparse distinct enlarged tubercles on the flanks; (14) single subgular vocal sac in males; (15) presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males; (16) dense tubercles on skin of upper lip, upper eyelid, mandibular articulation, loreal and temporal regions; (17) dense tubercles on skin of ventral surface of thigh, spiny tubercles surrounding the cloaca. Comparisons. Comparative data of Boulenophrys yunkaiensis sp. nov. from B. yaoshanensis sp. nov., B. yingdeensis sp. nov., and the other recognized members of the genus Boulenophrys are listed in Table 3. Having a smaller body size with SVL 35.3–40.0 mm in males, Boulenophrys yunkaiensis sp. nov. is significantly different from congeners whose SVL> 50 mm in males, including B. caudoprocta (81.3 mm in single male), B. jingdongensis (53.0– 56.5 mm in males), B. liboensis (60.5–67.7 mm in males), B. mirabilis (55.8–61.4 mm in males), B. omeimontis (56.0– 59.5 mm in males), B. sangzhiensis (54.7 mm in single male), and B. shuichengensis (102.0– 118.3 mm in males). Having relatively longer shanks with the heels overlapping or meeting when the flexed hindlimbs are held at right angles to the body axis, Boulenophrys yunkaiensis sp. nov. can be easily distinguished from the following nine congeners, B. acuta, B. brachykolos, B. daoji, B. dongguanensis, B. insularis, B. nankunensis, B. obesa, B. ombrophila, and B. wugongensis (vs. all of which have relatively shorter shanks with the heels not meeting). Lacking vomerine teeth, Boulenophrys yunkaiensis sp. nov. differs from B. caudoprocta, B. daiyunensis, B. daweimontis, B. dongguanensis, B. fansipanensis, B. frigida, B. hoanglienensis, B. insularis, B. jingdongensis, B. jinggangensis, B. jiulianensis, B. liboensis, B. nankunensis, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. rubrimera, B. sangzhiensis, B. shimentaina, B. tongboensis, and B. yingdeensis sp. nov. (vs. presence of vomerine teeth in these species). Having an unnotched tongue, Boulenophrys yunkaiensis sp. nov. can be distinguished from B. baolongensis, B. binlingensis, B. boettgeri, B. cheni, B. hoanglienensis, B. huangshanensis, B. insularis, B. jingdongensis, B. jiulianensis, B. kuatunensis, B. liboensis, B. lushuiensis, B. minor, B. nanlingensis, B. ombrophila, B. qianbeiensis, B. sangzhiensis, B. sanmingensis, B. shuichengensis B. spinata, and B. tongboensis (vs. tongue notched posteriorly in these species). Lacking lateral fringes on toes, Boulenophrys yunkaiensis sp. nov. differs from B. acuta, B. anlongensis, B. baishanzuensis, B. binchuanensis, B. boettgeri, B. congjiangensis, B. cheni, B. daiyunensis, B. daoji, B. jingdongensis, B. jinggangensis, B. liboensis, B. lini, B. lushuiensis, B. mirabilis, B. mufumontana, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. rubrimera, B. sangzhiensis, B. sanmingensis, B. shimentaina, B. shuichengensis, B. spinata, B. xiangnanensis, B. xianjuensis, and B. yangmingensis (vs. presence of lateral fringes on toes in these species); and from B. wushanensis (vs. presence of wide lateral fringes on toes in males while lacking in females). Having rudimentary webbing on toes, Boulenophrys yunkaiensis sp. nov. differs from B. baishanzuensis, B. baolongensis, B. daweimontis, B. fansipanensis, B. frigida, B. huangshanensis, B. kuatunensis, B. lishuiensis, B. ombrophila, B. rubrimera, B. tongboensis, and B. wuliangshanensis (vs. absence of webbing on toes in these species); and from B. jingdongensis, B. palpebralespinosa, B. qianbeiensis, B. shuichengensis, and B. spinata (vs. presence of well-developed webbing on toes in these species). For the remaining eight species, Boulenophrys yunkaiensis sp. nov. can be further distinguished by body size with SVL 45.3–46.1 mm in females (vs. 37.5–39.2 mm in B. angka, 39.5–40.4 mm in B. jiangi, 42.3 mm in B. leishanensis, 37.6 mm in B. shunhuangensis, and 50.5 mm in B. tuberogranulatus), the relative finger lengths II B. angka, IV B. caobangensis), and the presence of tiny spines on skin of upper lip, upper eyelid, loreal and temporal regions excluding the tympanum in adult males (vs. absence of such tiny spines in B. angka, B. chishuiensis, B. jiangi, B. leishanensis, B. shunhuangensis, B. tuberogranulatus, and B. yaoshanensis sp. nov.). Description of holotype. Adult male. small body size, SVL 37.0 mm; head width slightly larger than head length, HWD/HDL 1.15; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond the margin of lower jaw; top of head flat; eyes moderate in size, ED 0.36 of HDL, pupil vertical, near diamond-shaped; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanum boundary clear, TD/ED 1.84; large ovoid choanae at the base of the maxilla; vomerine ridge weak, vomerine teeth absent, maxillary teeth present; margin of tongue rounded, not notched behind; presence of a single subgular vocal sac, a pair of slit-like openings at posterior of jaw. Radio-ulnar length 0.22 of SVL and hand 0.27 of SVL; hand without webbing, fingers without lateral fringes, relative finger length II Skin of doesum rough with sparse granules; sparse large tubercles on the flanks; a small, horn-like prominent tubercle on the edge of upper eyelid; clear supratympanic fold curving posteroventrally from posterior corner of eye to a level above insertion of arm; dense tubercles on skin of upper lip, upper eyelid, mandibular articulation, loreal and temporal regions, excluding the tympanum and surface around cloaca, some of them bearing tiny spines; a discontinuous “X” shaped ridge and a few elongated warts arranged in longitudinal rows on two sides at the middorsum; sparse tubercles on the dorsal shank and thigh; ventral surface smooth; small pectoral gland close to axilla; a single femoral gland positioned subequally distant from knees and cloaca on posterior surface of each thigh. Coloration of holotype in life. Dorsal surface of body yellowish brown with an inverted brown triangular marking between eyes; an “X” shaped marking on the mid-dorsum. Forearms and hindlimbs with dark brown transverse bands. Supratympanic fold with a discontinuous white line; a dark vertical band below the eye, from the inferior margin of the eye to the upper lip. Numerous brown patches scattered on lateroventral surface of flanks; groin red-orange. Ventral surface of throat and chest light salmon in color with brown patches, an indistinct longitudinal stripe on throat; ventral surface of body light salmon in color with brown patches and white spots; ventral surface of limbs light salmon in color with dark brown spots and blotches; ventral surfaces of hands and ventral surfaces of feet brown, tips of digits pale brown; metacarpal tubercle and metatarsal tubercle reddish. Pectoral glands and femoral glands white. Iris yellowish brown. Coloration of holotype in preservative. Yellowish brown faded to greyish brown dorsally. Triangular marking between eyes, “X”-shaped marking on the mid-dorsum, transverse bands on dorsal forearms and hind limbs became indistinct. Color of ventral surface faded to greyish white, all bands and spots became indistinct. Variation and sexual dimorphism. Measurement data of type series are listed in Table 6. Females (SVL 45.3–46.1 mm) are distinctly larger than males (SVL 35.3 –40.0 mm). In adult males, skin of upper lip, upper eyelid, mandibular articulation, loreal and temporal regions excluding the tympanum and surface around cloaca with dense tubercles, some of them bearing tiny spine. Tubercles present in females but without tiny spines. Presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males. Distribution and ecology. Currently, Boulenophrys yunkaiensis sp. nov. is known only from Yunkaishan Nature Reserve, Guangdong, China.All individuals were found in evergreen secondary forest, near montane streams and in the nearby leaf litter on the forest floor at elevations between 900–1400 m. Males call on the rocks by the flowing streams from April to June, suggesting their breeding season corresponds to this period. Females were found on the forest floor, and one female was observed feeding on an earthworm. Tadpoles could be found all year-round.

I. Genus Megophrys Kuhl & Van Hasselt, 1822 Suggested common name: Indonesian horned toads (in English) / fflDzfl (in Chinese) Five species: Megophrys montana Kuhl & Van Hasselt, 1822 (type species); Megophrys acehensis Munir, Nishikawa, Hamidy & Smith, 2021; Megophrys lancip Munir, Hamidy, Farajallah & Smith, 2018; Megophrys parallela Inger & Iskandar, 2005; Megophrys selatanensis Munir, Nishikawa, Hamidy & Smith, 2021., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

VI. Genus Ophryophryne Boulenger, 1903 Suggested common name: Narrow-mouth horned toads (in English) / ���fflDzfl (in Chinese) Seven species: Ophryophryne microstoma Boulenger, 1903 (type species); Ophryophryne elfina Poyarkov, Duong, Orlov, Gogoleva, Vassilieva, Nguyen, Nguyen, Nguyen, Che & Mahony, 2017; Ophryophryne gerti Ohler, 2003; Ophryophryne hansi Ohler, 2003; Ophryophryne pachyproctus Kou, 1985; Ophryophryne poilani Bourret, 1937; Ophryophryne synoria Stuart, Sok & Neang, 2006., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on pages 430-431, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

2) Xenophrys aceras group Two species: Xenophrys aceras Boulenger, 1903; Xenophrys longipes (Boulenger, 1886)., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

Boulenophrys yunkaiensis sp. nov. Qi, Wang, Lyu & Wang Yunkai Horned Toad / yun kai jiao chan (������fflDz) Figures 6, 7C Chresonymy. Megophrys sp 32 (SYS a004637���4638, 4694)��� Liu et al. 2018 Holotype. SYS a004637 (Figs. 6A, 7C), adult male, collected on 14 April 2016 by Jian Wang, Zhi-Tong Lyu and Ying-Yong Wang from the Yunkaishan Nature Reserve (22.2758��N, 111.1952��E; ca 950 m a.s.l.), Xinyi City, Guangdong Province, China. Paratypes. Eight adult specimens from the same locality as the holotype: males SYS a004636 and SYS a004638 collected on 14 April 2016 by Jian Wang, Zhi-Tong Lyu and Ying-Yong Wang; male SYS a004662/ CIB 116085 and females SYS a004659 (Figs. 6B) and SYS a004660 collected on 15 April 2016 by Jian Wang, Zhi-Tong Lyu and Ying-Yong Wang; female SYS a004691 collected on 16 April 2016 by Jian Wang, Zhi-Tong Lyu and Ying-Yong Wang; males SYS a004986 and SYS a004987 collected on 28 June 2016 by Jian Wang. Etymology: The specific epithet yunkaiensis refers to its type locality, the Yunkaishan Nature Reserve, western Guangdong, China. Diagnosis. (1) Small body size, SVL 35.3���40.0 mm (37.6 �� 1.7, N = 6) in adult males and SVL 45.3���46.1 mm (45.8 �� 0.4, N = 3) in adult females; (2) snout rounded in dorsal view; (3) tympanum boundary clear, ED/TD 1.68���1.91 in males, 1.47���1.80 in females; (4) presence of vomerine ridge and absence of vomerine teeth; (5) margin of tongue rounded, not notched behind; (6) hindlimbs slender, heels overlapping or just meeting and tibio-tarsal articulation reaching forward between tympanum to posterior corner of eye when leg stretched forward; (7) tibia 0.40���0.48 of SVL and foot 0.60���0.68 of SVL in males, while tibia 0.42���0.46 of SVL and foot 0.59���0.66 of SVL in female; (8) a subarticular tubercle present at the base of each fingers; (9) toes without lateral fringes and with only rudimentary webbing; (10) presence of small horn-like tubercle at the edge of upper eyelid; (11) surface around cloaca with large tubercles bearing tiny spines; (12) dorsal skin rough with small granules, a discontinuous ���X��� or ���Y���-shaped ridge with two discontinuous dorsolateral ridges on two side of dorsum; (13) sparse distinct enlarged tubercles on the flanks; (14) single subgular vocal sac in males; (15) presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males; (16) dense tubercles on skin of upper lip, upper eyelid, mandibular articulation, loreal and temporal regions; (17) dense tubercles on skin of ventral surface of thigh, spiny tubercles surrounding the cloaca. Comparisons. Comparative data of Boulenophrys yunkaiensis sp. nov. from B. yaoshanensis sp. nov., B. yingdeensis sp. nov., and the other recognized members of the genus Boulenophrys are listed in Table 3. Having a smaller body size with SVL 35.3���40.0 mm in males, Boulenophrys yunkaiensis sp. nov. is significantly different from congeners whose SVL> 50 mm in males, including B. caudoprocta (81.3 mm in single male), B. jingdongensis (53.0��� 56.5 mm in males), B. liboensis (60.5���67.7 mm in males), B. mirabilis (55.8���61.4 mm in males), B. omeimontis (56.0��� 59.5 mm in males), B. sangzhiensis (54.7 mm in single male), and B. shuichengensis (102.0��� 118.3 mm in males). Having relatively longer shanks with the heels overlapping or meeting when the flexed hindlimbs are held at right angles to the body axis, Boulenophrys yunkaiensis sp. nov. can be easily distinguished from the following nine congeners, B. acuta, B. brachykolos, B. daoji, B. dongguanensis, B. insularis, B. nankunensis, B. obesa, B. ombrophila, and B. wugongensis (vs. all of which have relatively shorter shanks with the heels not meeting). Lacking vomerine teeth, Boulenophrys yunkaiensis sp. nov. differs from B. caudoprocta, B. daiyunensis, B. daweimontis, B. dongguanensis, B. fansipanensis, B. frigida, B. hoanglienensis, B. insularis, B. jingdongensis, B. jinggangensis, B. jiulianensis, B. liboensis, B. nankunensis, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. rubrimera, B. sangzhiensis, B. shimentaina, B. tongboensis, and B. yingdeensis sp. nov. (vs. presence of vomerine teeth in these species). Having an unnotched tongue, Boulenophrys yunkaiensis sp. nov. can be distinguished from B. baolongensis, B. binlingensis, B. boettgeri, B. cheni, B. hoanglienensis, B. huangshanensis, B. insularis, B. jingdongensis, B. jiulianensis, B. kuatunensis, B. liboensis, B. lushuiensis, B. minor, B. nanlingensis, B. ombrophila, B. qianbeiensis, B. sangzhiensis, B. sanmingensis, B. shuichengensis B. spinata, and B. tongboensis (vs. tongue notched posteriorly in these species). Lacking lateral fringes on toes, Boulenophrys yunkaiensis sp. nov. differs from B. acuta, B. anlongensis, B. baishanzuensis, B. binchuanensis, B. boettgeri, B. congjiangensis, B. cheni, B. daiyunensis, B. daoji, B. jingdongensis, B. jinggangensis, B. liboensis, B. lini, B. lushuiensis, B. mirabilis, B. mufumontana, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. rubrimera, B. sangzhiensis, B. sanmingensis, B. shimentaina, B. shuichengensis, B. spinata, B. xiangnanensis, B. xianjuensis, and B. yangmingensis (vs. presence of lateral fringes on toes in these species); and from B. wushanensis (vs. presence of wide lateral fringes on toes in males while lacking in females). Having rudimentary webbing on toes, Boulenophrys yunkaiensis sp. nov. differs from B. baishanzuensis, B. baolongensis, B. daweimontis, B. fansipanensis, B. frigida, B. huangshanensis, B. kuatunensis, B. lishuiensis, B. ombrophila, B. rubrimera, B. tongboensis, and B. wuliangshanensis (vs. absence of webbing on toes in these species); and from B. jingdongensis, B. palpebralespinosa, B. qianbeiensis, B. shuichengensis, and B. spinata (vs. presence of well-developed webbing on toes in these species). For the remaining eight species, Boulenophrys yunkaiensis sp. nov. can be further distinguished by body size with SVL 45.3���46.1 mm in females (vs. 37.5���39.2 mm in B. angka, 39.5���40.4 mm in B. jiangi, 42.3 mm in B. leishanensis, 37.6 mm in B. shunhuangensis, and 50.5 mm in B. tuberogranulatus), the relative finger lengths II B. angka, IV B. caobangensis), and the presence of tiny spines on skin of upper lip, upper eyelid, loreal and temporal regions excluding the tympanum in adult males (vs. absence of such tiny spines in B. angka, B. chishuiensis, B. jiangi, B. leishanensis, B. shunhuangensis, B. tuberogranulatus, and B. yaoshanensis sp. nov.). Description of holotype. Adult male. small body size, SVL 37.0 mm; head width slightly larger than head length, HWD/HDL 1.15; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond the margin of lower jaw; top of head flat; eyes moderate in size, ED 0.36 of HDL, pupil vertical, near diamond-shaped; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanum boundary clear, TD/ED 1.84; large ovoid choanae at the base of the maxilla; vomerine ridge weak, vomerine teeth absent, maxillary teeth present; margin of tongue rounded, not notched behind; presence of a single subgular vocal sac, a pair of slit-like openings at posterior of jaw. Radio-ulnar length 0.22 of SVL and hand 0.27 of SVL; hand without webbing, fingers without lateral fringes, relative finger length II Coloration of holotype in life. Dorsal surface of body yellowish brown with an inverted brown triangular marking between eyes; an ���X��� shaped marking on the mid-dorsum. Forearms and hindlimbs with dark brown transverse bands. Supratympanic fold with a discontinuous white line; a dark vertical band below the eye, from the inferior margin of the eye to the upper lip. Numerous brown patches scattered on lateroventral surface of flanks; groin red-orange. Ventral surface of throat and chest light salmon in color with brown patches, an indistinct longitudinal stripe on throat; ventral surface of body light salmon in color with brown patches and white spots; ventral surface of limbs light salmon in color with dark brown spots and blotches; ventral surfaces of hands and ventral surfaces of feet brown, tips of digits pale brown; metacarpal tubercle and metatarsal tubercle reddish. Pectoral glands and femoral glands white. Iris yellowish brown. Coloration of holotype in preservative. Yellowish brown faded to greyish brown dorsally. Triangular marking between eyes, ���X���-shaped marking on the mid-dorsum, transverse bands on dorsal forearms and hind limbs became indistinct. Color of ventral surface faded to greyish white, all bands and spots became indistinct. Variation and sexual dimorphism. Measurement data of type series are listed in Table 6. Females (SVL 45.3���46.1 mm) are distinctly larger than males (SVL 35.3 ���40.0 mm). In adult males, skin of upper lip, upper eyelid, mandibular articulation, loreal and temporal regions excluding the tympanum and surface around cloaca with dense tubercles, some of them bearing tiny spine. Tubercles present in females but without tiny spines. Presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males. Distribution and ecology. Currently, Boulenophrys yunkaiensis sp. nov. is known only from Yunkaishan Nature Reserve, Guangdong, China.All individuals were found in evergreen secondary forest, near montane streams and in the nearby leaf litter on the forest floor at elevations between 900���1400 m. Males call on the rocks by the flowing streams from April to June, suggesting their breeding season corresponds to this period. Females were found on the forest floor, and one female was observed feeding on an earthworm. Tadpoles could be found all year-round., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on pages 421-424, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979, {"references":["Liu, Z. Y., Chen, G. L., Zhu, T. Q., Zeng, Z. C., Lyu, Z. T., Wang, J., Messenger, K., Greenberg, A. J., Guo, Z. X., Yang, Z. H., Shi, S. H. & Wang, Y. Y. (2018) Prevalence of cryptic species in morphologically uniform taxa - Fast speciation and evolutionary radiation in Asian frogs. Molecular Phylogenetics and Evolution, 127, 723 - 731. https: // doi. org / 10.1016 / j. ympev. 2018.06.020"]}

1) Xenophrys major group Twelve species: Xenophrys monticola G��nther, 1864 (type species); Xenophrys major (Boulenger, 1908); Xenophrys flavipunctata (Mahony, Kamei, Teeling & Biju, 2018); Xenophrys glandulosa (Fei, Ye & Huang, 1990); Xenophrys himalayana (Mahony, Kamei, Teeling & Biju, 2018); Xenophrys maosonensis (Bourret, 1937); Xenophrys mangshanensis (Fei & Ye, 1990); Xenophrys medogensis (Fei, Ye & Huang, 1983); Xenophrys oreocrypta (Mahony, Kamei, Teeling & Biju, 2018); Xenophrys periosa (Mahony, Kamei, Teeling & Biju, 2018); Xenophrys robusta (Boulenger, 1908); Xenophrys zhangi (Ye & Fei, 1992)., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

Boulenophrys yaoshanensis sp. nov. Qi, Mo, Lyu, Wang & Wang Mt. Dayao Horned Toad / yao shan jiao chan (Ṉ山fflDz) Figures 3, 7A Chresonymy. Megophrys minor — Liu & Hu 1962 Megophrys brachykolos — Fei et al. 2009; Mo et al. 2014 Boulenophrys brachykolos — Fei & Ye 2016; Fei 2020 Megophrys sp 31 (SYS a002189–2190, 4850–4851, 4878)— Liu et al. 2018 Holotype. SYS a002189 (Figures 3A, 7A), adult male, collected on 8 July 2013 by Yu-Long Li and Ying-Yong Wang from the Dayaoshan Nature Reserve (26.5517°N, 114.1548°E; ca 845 m a.s.l.), Jinxiu Yao Autonomous County, Guangxi Zhuang Autonomous Region, China. Paratypes. Five adult specimens from the same locality as the holotype: male SYS a000838 collected on 13 September 2011 by Yu-Long Li and Ying-Yong Wang; males SYS a004850 and SYS a004851 / CIB 116086 collected on 1 June 2016 by Jian Wang; female SYS a004878 (Figure 3B) collected on 3 June 2016 by Jian Wang; male SYS a007023 collected on 1 June 2019 by Zhi-Tong Lyu and Yu-Long Li. Female NHMG1503016 collected on 18 March 2015 by Yun-Ming Mo from the Mt. Xianglu (24.1055°N, 110.2300°E; ca 1305 m a.s.l.), Jinxiu Yao Autonomous County; male NHMG201705032 collected on 5 May 2017 by Yun-Ming Mo from 16 km west of Jinxiu Yao Autonomous County (24.1162°N, 110.2491 °E, ca 1115m a.s.l.), Guangxi Zhuang Autonomous Region, China. Etymology. The specific epithet yaoshanensis refers to the type locality, the Dayaoshan Nature Reserve, Guangxi, China. Particularly, we employ the epithet “yaoshan” rather than “dayaoshan” to make it consistent with other zoological and botanical species discovered from this area, e.g. Zhangixalus yaoshanensis (Liu & Hu, 1962), Leptobrachium liui yaoshanensis (Liu & Hu, 1978), Litsea yaoshanensis Yang & Huang, 1978, and Rhododendron yaoshanicum Fang & He, 1983 (Liu & Hu, 1962; Liu et al. 1978; Yang et al. 1978; Fang & He, 1983). Diagnosis. (1) Small body size, SVL 32.5–42.6 mm (37.1 ± 3.5, N = 6) in adult males and SVL 46.6–47.4 mm (47.0 ± 0.6, N = 2 in adult females; (2) snout rounded in dorsal view; (3) tympanum boundary clear, ED/TD 1.38–1.85 in males, 1.50–1.77 in females; (4) weak vomerine ridge present, vomerine teeth absent; (5) margin of tongue rounded, not notched posteriorly; (6) hind limbs slender, heels slightly overlapping or meeting, and tibiotarsal articulation reaching eye when leg stretched forward; (7) tibia 0.45–0.51 of SVL and foot 0.63–0.73 of SVL in males, tibia 0.44–0.47 of SVL and foot 0.63–0.67 of SVL in females; (8) toes without lateral fringes and with rudimentary webbing; (9) small horn-like tubercle at the edge of upper eyelid present; (10) skin of dorsum relatively smooth, a discontinuous “X”-shaped ridge on at mid-dorsum; (11) a few sparsely distributed large tubercles on flanks; (12) body yellowish brown dorsally, an inverted hollow dark-brown triangularly shaped marking between eyes, a dark “X”-shaped making at mid-dorsum; (13) single subgular vocal sac in males; (14) presence of villiform black nuptial spines on dorsal surface of first and second fingers in adult males. Comparisons. Comparative data of Boulenophrys yaoshanensis sp. nov. to other recognized members of the genus Boulenophrys are listed in Table 3. Having a smaller body size with SVL 32.5–42.6 mm in males, Boulenophrys yaoshanensis sp. nov. differs from the seven congeners whose SVL> 50 mm in males, including B. caudoprocta (81.3 mm in a single male), B. jingdongensis (53.0– 56.5 mm), B. liboensis (60.5–67.7 mm), B. mirabilis (55.8–61.4 mm), B. omeimontis (56.0– 59.5 mm), B. sangzhiensis (54.7 mm in a single male), and B. shuichengensis (102.0– 118.3 mm). Having relatively longer shanks with heels that overlap or meet when the flexed hind limbs are held at right angles to the body axis, Boulenophrys yaoshanensis sp. nov. can be easily distinguished from the following nine congeners, B. acuta, B. brachykolos, B. daoji, B. dongguanensis, B. insularis, B. nankunensis, B. obesa, B. ombrophila, and B. wugongensis (vs. all of which have relatively shorter shanks with the heels not meeting). Lacking vomerine teeth, Boulenophrys yaoshanensis sp. nov. differs from B. caudoprocta, B. daiyunensis, B. daweimontis, B. dongguanensis, B. fansipanensis, B. frigida, B. hoanglienensis, B. insularis, B. jingdongensis, B. jinggangensis, B. jiulianensis, B. liboensis, B. nankunensis, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. rubrimera, B. sangzhiensis, B. shimentaina, and B. tongboensis (vs. presence of vomerine teeth in these species). Having an unnotched tongue, Boulenophrys yaoshanensis sp. nov. can be distinguished from B. baolongensis, B. binlingensis, B. boettgeri, B. cheni, B. hoanglienensis, B. huangshanensis, B. insularis, B. jingdongensis, B. jiulianensis, B. kuatunensis, B. liboensis, B. lushuiensis, B. minor, B. nanlingensis, B. ombrophila, B. qianbeiensis, B. sangzhiensis, B. sanmingensis, B. shuichengensis B. spinata, and B. tongboensis (vs. tongue notched posteriorly in these species). Lacking lateral fringes on toes, Boulenophrys yaoshanensis sp. nov. differs from B. acuta, B. anlongensis, B. baishanzuensis, B. binchuanensis, B. boettgeri, B. congjiangensis, B. cheni, B. daiyunensis, B. daoji, B. jingdongensis, B. jinggangensis, B. liboensis, B. lini, B. lushuiensis, B. mirabilis, B. mufumontana, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. rubrimera, B. sangzhiensis, B. sanmingensis, B. shimentaina, B. shuichengensis, B. spinata, B. xiangnanensis, B. xianjuensis, and B. yangmingensis (vs. presence of lateral fringes on toes in these species); and from B. wushanensis (vs. presence of wide lateral fringes on toes in males while lacking in females). Having rudimentary webbing on toes, Boulenophrys yaoshanensis sp. nov. differs from B. baishanzuensis, B. baolongensis, B. daweimontis, B. fansipanensis, B. frigida, B. huangshanensis, B. kuatunensis, B. lishuiensis, B. ombrophila, B. rubrimera, B. tongboensis, and B. wuliangshanensis (vs. absence of webbing on toes in these species); and from B. jingdongensis, B. palpebralespinosa, B. qianbeiensis, B. shuichengensis, and B. spinata (vs. presence of well-developed webbing on toes in these species). For the remaining seven species, Boulenophrys yaoshanensis sp. nov. can be further distinguished by the body size with SVL 46.6–47.4 mm in females (vs. 37.5–39.2 mm in B. angka, 39.5–40.4 mm in B. jiangi, 42.3 mm in B. leishanensis, 37.6 mm in B. shunhuangensis, and 50.5 mm in B. tuberogranulatus), the relative finger lengths II B. angka, IV B. caobangensis), tibio-tarsal articulation reaching eye when leg stretched forward (vs. between the nasal and tip of snout in B. shunhuangensis; between tympanum and eye in B. chishuiensis), dorsal skin relatively smooth (vs. dorsal skin rough with numerous granules in B. chishuiensis and B. tuberogranulatus), and absence of any spines on head (vs. presence of small black spines on temporal region in B. caobangensis). Description of holotype. Adult male. small body size, SVL 37.3 mm; head length equal to head width, HDW/ HDL 1.00; snout rounded in dorsal view, projecting well beyond the margin of lower jaw, sloping backward to mouth lateral in profile; top of head flat; eyes moderate in size, ED 0.38 of HDL, pupil vertical, near diamondshaped; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanum clear, ED/TD 1.68; large ovoid choanae at the base of the maxilla; weak vomerine ridge present, vomerine teeth absent, maxillary teeth present; margin of tongue rounded, not notched posteriorly; a single subgular vocal sac present, and a pair of slit-like openings at posterior of jaw. Radio-ulnar length 0.23 of SVL and hand 0.27 of SVL; hand without webbing, fingers without lateral fringes, relative finger length II Skin of dorsum relatively smooth with small granules; sparsely distributed large tubercles on flanks; a small, horn-like prominent tubercle on the edge of upper eyelid; unpigmented supratympanic fold curving posteroventrally from posterior margin of eye to a level above insertion of arm; a discontinuous “X”-shaped ridge at mid-dorsum; a few sparsely distributed large tubercles on flanks; sparsely distributed tubercles on dorsal surface of foreleg and thigh; ventral surfaces smooth; area around cloaca bearing small, spinose tubercles males; small pectoral gland close to axillae; and a single femoral gland on positioned subequally distant from knees and cloaca on posterior surface of each thigh.. Coloration of holotype in life. Dorsal surface of body yellowish brown with an inverted hollow dark-brown triangularly shaped marking between eyes; an “X” shaped marking on the mid-dorsum. Forearms and hind limbs with dark-brown transverse bands. Supratympanic fold light-colored; a dark vertical band below the eye, from the inferior margin of the eye to the upper lip. Ventral surface of throat and chest greyish brown with dark-brown patches; ventral surface of body greyish white with creamy white and orange spots, ventral surface of limbs greyish brown with dark brown spots; ventral surfaces of hands and feet brown, tips of digits pale brown; metacarpal tubercle and metatarsal tubercle reddish. Pectoral gland and femoral gland white. Iris yellowish brown. Coloration of holotype in preservative. Yellowish brown faded to greyish brown dorsally. Triangular marking between eyes, “X” shaped marking on the mid-dorsum, transverse bands on dorsal forearms and hind limbs became indistinct. Color of ventral surface faded to greyish white, all bands and spots became indistinct. Variation and sexual dimorphism. Mensural data of the type series are listed in Table 4. Two females (SYS a004878, SVL 46.6 mm; NHMG1503016, SVL 47.4 mm) are distinctly larger than the males (SVL 32.5–42.6 mm). Three paratypes (SYS a004850–4851, 4878) have subarticular tubercles on the first phalangeal articulations of the fingers. The holotype and two male paratypes (SYS a004850–4851) have small, spinose tubercles around the cloaca. Villiform black nuptial spines occur on the dorsal surface of the first and second fingers in adult males. Distribution and ecology. Currently, Boulenophrys yaoshanensis sp. nov. is known only from Jinxiu and Mengshan counties, Guangxi, China. All individuals were found in evergreen secondary forest, inhabiting flowing montane streams and the nearby forest floor and leaf litter at elevations between 800–1350 m. Males call while perched on leaves from May to July, suggesting their breeding season corresponds to this period. Females were found on the forest floor and tadpoles were not observed.

2) Brachytarsophrys orientalis group Five species: Brachytarsophrys orientalis Li, Lyu, Wang & Wang, 2020; Brachytarsophrys chuannanensis Fei, Ye & Huang, 2001; Brachytarsophrys feae (Boulenger, 1887); Brachytarsophrys platyparietus Rao & Yang, 1997; Brachytarsophrys popei Zhao, Yang, Chen, Chen & Wang, 2014., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

Specimens of genus Boulenophrys examined in this study. B. acuta (10): China: Guangdong: Fengkai: Heishiding Nature Reserve (type locality): SYS a000168, 0517, 0521, 3257, 2159, 2266���2269, 2276. B. binlingensis (2): China: Sichuan: Hongya: Mt. Wawu (type locality): SYS a005313���5314. B. boettgeri (16): China: Fujian: Wuyishan: Mt. Wuyi (type locality): SYS a002480, 4149���4151; Jiangxi: Guixi: Yangjifeng Nature Reserve: SYS a000312, 0315, 0328���0330, 0376, 0378; Guangfeng: Tongboshan Nature Reserve: SYS a001671��� 1673, 1683, 1700. B. brachykolos (12): China: Hong Kong (type locality): SYS a001502���1503; Guangdong: Shenzhen: Mt. Yangtai: SYS a002051���56; Dapeng Peninsula: SYS a002406���2408, 2410. B. caudoprocta (3): China: Hunan: Sangzhi: Badagongshan Nature Reserve (type locality): SYS a004281, 4308���4309. B. cheni (19): China: Jiangxi: Jinggangshan: Mt. Jinggang (type locality): SYS a001427���1429, 1871���1873; Hunan: Yanling: Taoyuandong Nature Reserve: SYS a002123���2127, 2140���2145. B. daiyunensis (7): China: Fujian: Quanzhou: Daiyun Village (type locality): SYS a001730���1733, 6002; Jiuxianshan: SYS a006000, 6003. B. daoji (6): China: Zhejiang: Taizhou: Mt Tiantai (type locality): SYS a006209���6214 B. huangshanensis (13): China: Anhui: Huangshan: Mt. Huangshan (type locality): SYS a002702���2707; Jiangxi: Wuyuan: Mt. Dazhang: SYS a001622���1623, 3705���3707; Zhejiang: Lin���an: Mt. Tianmu: SYS a002684���2685. B. jingdongensis (24): China: Yunnan: Jingdong: Mt. Wuliang (type locality): SYS a003909, 3928���3929; Zhenyuan: Mt. Ailao: SYS a001778, 2988, 2989���2991, 2993���2994, 3005���3006, 3903���3904; Guangxi: Tianlin: Mt. Cenwanglao: SYS a005160���5165, 5184, 5968���5970. B. jinggangensis (11): China: Jiangxi: Jinggangshan: Mt. Jinggang (type locality): SYS a001413���1416, 1430, 4028; Hunan: Yanling: Taoyuandong Nature Reserve: SYS a001859���1863. B. kuatunensis (3): China: Fujian: Wuyishan: Guadun: SYS a001579, 1590; Jiangxi: Guixi: Yangjifeng Nature Reserve: SYS a000241. B. lini (27): China: Hunan: Yanling: Taoyuandong Nature Reserve (type locality): SYS a002128; Jiangxi: Jinggangshan: Mt. Jinggang: SYS a001417��� 1424, 2375���2386; Suichuan: Nanfengmian Nature Reserve: SYS a002369���2374. B. minor (5): China: Sichuan: Dujiangyan: Mt. Qingcheng (type locality): SYS a003209���3213. B. mirabilis (4): China: Guangxi: Longsheng: Huaping Nature Reserve (type locality): SYS a002192���2193, 2289, 2917. B. obesa (4): China: Guangdong: Fengkai: Heishiding Nature Reserve (type locality): SYS a002270���2272, 3047. B. omeimontis (11): China: Sichuan: Emeishan: Mt. Emei (type locality): SYS a001798���1801, 1940���1941, 5301; Hongya: Mt. Wawu: SYS a005330���5331; Pingshan: Mt. Laojun: SYS a002740���2741. B. sangzhiensis (6): China: Hunan: Sangzhi: Badagongshan Nature Reserve (type locality): SYS a004306���4307, 4313���4316. B. sanmingensis (7): China: Fujian: Sanming: Mt. Longtou (type locality): SYS a002493���2496, 2498���2500. B. shimentaina (11): China: Guangdong: Yingde: Shimentai Nature Reserve (type locality): SYS a002077, 2081���2085, 4172��� 4173, 4710, 5992���5993. B. spinata (2): China: Guizhou: Leishan: Mt. Leigong (type locality): SYS a002226���2227. B. tongboensis (5): China: Jiangxi: Shangrao: Mt. Tongbo (type locality): SYS a001911, 3225���3228. B. tuberogranulatus (1): China: Hunan: Sangzhi: Badagongshan Nature Reserve (type locality): SYS a004310. B. wushanensis (5): China: Hubei: Shennongjia: Shennongjia Nature Reserve (type locality): SYS a003008���3011, 3013. B. wuliangshanensis (5): China: Yunnan: Jingdong: Mt. Wuliang (type locality): SYS a003924���3925; Zhenyuan: Mt. Ailao: SYS a002983���29. B. xiangnanensis (11): China: Hunan: Shuangpai: Mt. Yangming (type locality): SYS a002874���2875, 2878���2886. B. yangmingensis (7): China: Hunan: Shuangpai: Mt. Yangming (type locality): SYS a002877, 2887���2889, 2891���2892., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on pages 431-432, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

Boulenophrys yingdeensis sp. nov. Qi, Lyu, Wang & Wang Yingde Horned Toad / ying de jiao chan (�����fflDz) Figures 4, 5, 7B Chresonymy. Megophrys sp 4 (SYS a002100, 4721, 5447)��� Liu et al. 2018 Holotype. SYS a002099 (Figures 4A, 7B), adult male, collected on 26 April 2013 by Run-Lin Li from Shimentai Nature Reserve (24.4435��N, 113.3034��E; ca 357 m a.s.l.), Yingde City, Guangdong Province, China. Paratypes. Eight adult specimens from the same locality as the holotype: female SYS a001563 collected on 23 April 2012 by Run-Lin Li; female SYS a004721 collected on 29 April 2016 by Ying-Yong Wang, Jian Wang and Zhi-Tong Lyu; female SYS a005447 and male SYS a005449 collected on 19 August 2016 by Zhi-Tong Lyu; male SYS a007114/ CIB 116084 collected on 20 June 2018 by Hong-Hui Chen, Jia-He Li and Jian Wang; male SYS a007405 and two females SYS a007406 and SYS a007407 collected on 20 August 2018 by Jian Wang. Etymology. The specific epithet yingdeensis refers to its type locality, Yingde City in northern Guangdong. Diagnosis. (1) Small body size, SVL 33.2���35.3 mm (34.2 �� 1.0, N = 4) in adult males and SVL 36.3���45.8 mm (40.7 �� 4.2, N = 5) in adult females; (2) snout rounded in dorsal view; (3) tympanum boundary clear, ED/TD 1.65��� 1.95 in males, 1.48���2.09 in females; (4) vomerine ridge prominent bearing vomerine teeth; (5) margin of tongue rounded, not notched behind; (6) hindlimbs slender, heels overlapping or just meeting and tibio-tarsal articulation reaching forward between tympanum to posterior corner of eye; (7) tibia 0.46���0.48 of SVL and foot 0.61���0.67 of SVL in males, while tibia 0.44���0.46 of SVL and foot 0.61���0.66 of SVL in females; (8) toes without lateral fringes and with only rudimentary webbing; (9) presence of small, horn-like tubercle at the edge of upper eyelid; (10) dorsal skin smooth, a discontinuous ���Y��� or ���X���-shaped ridge on the mid-dorsum, two discontinuous dorsolateral ridges on two side on the dorsum; (11) skin of flanks smooth with small conical tubercles; (12) single subgular vocal sac in males; (13) presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males. Comparisons. Comparative data of Boulenophrys yingdeensis sp. nov. from B. yaoshanensis sp. nov. and the other recognized members of the genus Boulenophrys are listed in Table 3. Having a smaller body size with SVL 33.2���35.3 mm in males, Boulenophrys yingdeensis sp. nov. differs from seven congeners whose SVL> 50 mm in males, including B. caudoprocta (81.3 mm in single male), B. jingdongensis (53.0��� 56.5 mm in males), B. liboensis (60.5���67.7 mm in males), B. mirabilis (55.8���61.4 mm in males), B. omeimontis (56.0��� 59.5 mm in males), B. sangzhiensis (54.7 mm in single male), and B. shuichengensis (102.0��� 118.3 mm in males). Having relatively longer shanks with the heels overlapping or meeting when the flexed hindlimbs are held at right angles to the body axis, Boulenophrys yingdeensis sp. nov. can be easily distinguished from the following nine congeners, B. acuta, B. brachykolos, B. daoji, B. dongguanensis, B. insularis, B. nankunensis, B. obesa, B. ombrophila, and B. wugongensis (vs. all of which have relatively shorter shanks with the heels not meeting). Having vomerine teeth, Boulenophrys yingdeensis sp. nov. differs from B. acuta, B. angka, B. anlongensis, B. baishanzuensis, B. baolongensis, B. binchuanensis, B. binlingensis, B. boettgeri, B. brachykolos, B. caobangensis, B. congjiangensis, B. cheni, B. chishuiensis, B. daoji, B. huangshanensis, B. jiangi, B. kuatunensis, B. leishanensis, B. lini, B. lishuiensis, B. lushuiensis, B. minor, B. mirabilis, B. mufumontana, B. obesa, B. ombrophila, B. sanmingensis, B. shuichengensis, B. shunhuangensis, B. spinata, B. tuberogranulatus, B. wugongensis, B. wuliangshanensis, B. wushanensis, B. xiangnanensis, B. xianjuensis, B. yangmingensis, and B. yaoshanensis sp. nov. (vs. absence of vomerine teeth in these species). Having an unnotched tongue, Boulenophrys yingdeensis sp. nov. can be distinguished from B. baolongensis, B. binlingensis, B. boettgeri, B. cheni, B. hoanglienensis, B. huangshanensis, B. insularis, B. jingdongensis, B. jiulianensis, B. kuatunensis, B. liboensis, B. lushuiensis, B. minor, B. nanlingensis, B. omeimontis, B. qianbeiensis, B. sangzhiensis, B. sanmingensis, B. shuichengensis B. spinata, and B. tongboensis (vs. tongue notched posteriorly in these species). Lacking lateral fringes on toes, Boulenophrys yingdeensis sp. nov. differs from B. acuta, B. anlongensis, B. baishanzuensis, B. binchuanensis, B. boettgeri, B. congjiangensis, B. cheni, B. daiyunensis, B. daoji, B. jingdongensis, B. jinggangensis, B. liboensis, B. lini, B. lushuiensis, B. mirabilis, B. mufumontana, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. rubrimera, B. sangzhiensis, B. sanmingensis, B. shimentaina, B. shuichengensis, B. spinata, B. xiangnanensis, B. xianjuensis, and B. yangmingensis (vs. presence of lateral fringes on toes in these species); and from B. wushanensis (vs. presence of wide lateral fringes on toes in males while lacking in females). Having rudimentary webbing on toes, Boulenophrys yingdeensis sp. nov. differs from B. baishanzuensis, B. baolongensis, B. daweimontis, B. fansipanensis, B. frigida, B. huangshanensis, B. kuatunensis, B. lishuiensis, B. ombrophila, B. rubrimera, B. tongboensis, and B. wuliangshanensis (vs. absence of webbing on toes in these species); and from B. jingdongensis, B. palpebralespinosa, B. qianbeiensis, B. shuichengensis, and B. spinata (vs. presence of well-developed webbing on toes in these species). Description of holotype. Adult male. small body size, SVL 35.3 mm; head width slightly larger than head length, HWD/HDL 1.07; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eyes moderate in size, ED 0.30 of HDL, pupil vertical, near diamond-shaped; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanum boundary clear, ED/TD 1.68; large ovoid choanae at the base of the maxilla; vomerine ridge prominent, vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched behind; presence of a single subgular vocal sac, a pair of slit-like openings at posterior of jaw. Radio-ulnar length 0.20 of SVL and hand 0.26 of SVL; hand without webbing, fingers without lateral fringes, relative finger length I Coloration of holotype in life. Dorsal surface of body olive-brown with an inverted brown triangular marking between eyes; an ���X��� shaped marking on the mid-dorsum. Forearms and hindlimbs with brown transverse bands. Supratympanic fold with a continuous white line; a dark vertical band below the eye, from the inferior margin of the eye to the upper lip. Ventral surface of throat and chest grayish brown with brown patches, an indistinct longitudinal stripe on surface of throat; a pair of dark brown longitudinal stripes scattered on surface of lateroventral flanks; ventral surface of body white with brown patches; ventral surface of limbs grayish brown with dark brown spots and blotches; ventral surfaces of hands and ventral surfaces of feet brown, tips of digits pale-brown; metacarpal tubercle and metatarsal tubercle reddish. Pectoral gland and femoral gland white. Iris yellowish brown. Coloration of holotype in preservative. Olive-brown faded to greyish brown dorsally. Triangular marking between eyes, ���X��� shaped marking on the mid-dorsum, transverse bands on dorsal forearms and hind limbs became indistinct. Color of ventral surface faded to greyish white all bands and spots became indistinct. Variation and sexual dimorphism. Measurement data of type series are listed in Table 5. Females (SVL 36.3���45.8 mm) are larger than males (SVL 33.2���35.3 mm). Presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males. Body coloration is quite variable, light brown, reddish brown, olive brown or dark brown dorsally (Figure 5). Distribution and ecology. Currently, Boulenophrys yingdeensis sp. nov. is known only from Shimentai Nature Reserve, Guangdong, China, and sympatric with B. shimentaina. All individuals were found in evergreen secondary forest, near lowland streams and nearbyleaf litter at elevations between 300��� 400 m. Males perch and call on plant leaves from April to June, suggesting their breeding season corresponds to this period. Females were found on the forest floor and tadpoles were not observed., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on pages 416-420, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979, {"references":["Liu, Z. Y., Chen, G. L., Zhu, T. Q., Zeng, Z. C., Lyu, Z. T., Wang, J., Messenger, K., Greenberg, A. J., Guo, Z. X., Yang, Z. H., Shi, S. H. & Wang, Y. Y. (2018) Prevalence of cryptic species in morphologically uniform taxa - Fast speciation and evolutionary radiation in Asian frogs. Molecular Phylogenetics and Evolution, 127, 723 - 731. https: // doi. org / 10.1016 / j. ympev. 2018.06.020"]}

V. Genus Xenophrys G��nther, 1864 Suggested common name: Strange horned toads (in English) / ���fflDzfl (in Chinese), Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

Boulenophrys yingdeensis sp. nov. Qi, Lyu, Wang & Wang Yingde Horned Toad / ying de jiao chan (Ȓ德fflDz) Figures 4, 5, 7B Chresonymy. Megophrys sp 4 (SYS a002100, 4721, 5447)— Liu et al. 2018 Holotype. SYS a002099 (Figures 4A, 7B), adult male, collected on 26 April 2013 by Run-Lin Li from Shimentai Nature Reserve (24.4435°N, 113.3034°E; ca 357 m a.s.l.), Yingde City, Guangdong Province, China. Paratypes. Eight adult specimens from the same locality as the holotype: female SYS a001563 collected on 23 April 2012 by Run-Lin Li; female SYS a004721 collected on 29 April 2016 by Ying-Yong Wang, Jian Wang and Zhi-Tong Lyu; female SYS a005447 and male SYS a005449 collected on 19 August 2016 by Zhi-Tong Lyu; male SYS a007114/ CIB 116084 collected on 20 June 2018 by Hong-Hui Chen, Jia-He Li and Jian Wang; male SYS a007405 and two females SYS a007406 and SYS a007407 collected on 20 August 2018 by Jian Wang. Etymology. The specific epithet yingdeensis refers to its type locality, Yingde City in northern Guangdong. Diagnosis. (1) Small body size, SVL 33.2–35.3 mm (34.2 ± 1.0, N = 4) in adult males and SVL 36.3–45.8 mm (40.7 ± 4.2, N = 5) in adult females; (2) snout rounded in dorsal view; (3) tympanum boundary clear, ED/TD 1.65– 1.95 in males, 1.48–2.09 in females; (4) vomerine ridge prominent bearing vomerine teeth; (5) margin of tongue rounded, not notched behind; (6) hindlimbs slender, heels overlapping or just meeting and tibio-tarsal articulation reaching forward between tympanum to posterior corner of eye; (7) tibia 0.46–0.48 of SVL and foot 0.61–0.67 of SVL in males, while tibia 0.44–0.46 of SVL and foot 0.61–0.66 of SVL in females; (8) toes without lateral fringes and with only rudimentary webbing; (9) presence of small, horn-like tubercle at the edge of upper eyelid; (10) dorsal skin smooth, a discontinuous “Y” or “X”-shaped ridge on the mid-dorsum, two discontinuous dorsolateral ridges on two side on the dorsum; (11) skin of flanks smooth with small conical tubercles; (12) single subgular vocal sac in males; (13) presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males. Comparisons. Comparative data of Boulenophrys yingdeensis sp. nov. from B. yaoshanensis sp. nov. and the other recognized members of the genus Boulenophrys are listed in Table 3. Having a smaller body size with SVL 33.2–35.3 mm in males, Boulenophrys yingdeensis sp. nov. differs from seven congeners whose SVL> 50 mm in males, including B. caudoprocta (81.3 mm in single male), B. jingdongensis (53.0– 56.5 mm in males), B. liboensis (60.5–67.7 mm in males), B. mirabilis (55.8–61.4 mm in males), B. omeimontis (56.0– 59.5 mm in males), B. sangzhiensis (54.7 mm in single male), and B. shuichengensis (102.0– 118.3 mm in males). Having relatively longer shanks with the heels overlapping or meeting when the flexed hindlimbs are held at right angles to the body axis, Boulenophrys yingdeensis sp. nov. can be easily distinguished from the following nine congeners, B. acuta, B. brachykolos, B. daoji, B. dongguanensis, B. insularis, B. nankunensis, B. obesa, B. ombrophila, and B. wugongensis (vs. all of which have relatively shorter shanks with the heels not meeting). Having vomerine teeth, Boulenophrys yingdeensis sp. nov. differs from B. acuta, B. angka, B. anlongensis, B. baishanzuensis, B. baolongensis, B. binchuanensis, B. binlingensis, B. boettgeri, B. brachykolos, B. caobangensis, B. congjiangensis, B. cheni, B. chishuiensis, B. daoji, B. huangshanensis, B. jiangi, B. kuatunensis, B. leishanensis, B. lini, B. lishuiensis, B. lushuiensis, B. minor, B. mirabilis, B. mufumontana, B. obesa, B. ombrophila, B. sanmingensis, B. shuichengensis, B. shunhuangensis, B. spinata, B. tuberogranulatus, B. wugongensis, B. wuliangshanensis, B. wushanensis, B. xiangnanensis, B. xianjuensis, B. yangmingensis, and B. yaoshanensis sp. nov. (vs. absence of vomerine teeth in these species). Having an unnotched tongue, Boulenophrys yingdeensis sp. nov. can be distinguished from B. baolongensis, B. binlingensis, B. boettgeri, B. cheni, B. hoanglienensis, B. huangshanensis, B. insularis, B. jingdongensis, B. jiulianensis, B. kuatunensis, B. liboensis, B. lushuiensis, B. minor, B. nanlingensis, B. omeimontis, B. qianbeiensis, B. sangzhiensis, B. sanmingensis, B. shuichengensis B. spinata, and B. tongboensis (vs. tongue notched posteriorly in these species). Lacking lateral fringes on toes, Boulenophrys yingdeensis sp. nov. differs from B. acuta, B. anlongensis, B. baishanzuensis, B. binchuanensis, B. boettgeri, B. congjiangensis, B. cheni, B. daiyunensis, B. daoji, B. jingdongensis, B. jinggangensis, B. liboensis, B. lini, B. lushuiensis, B. mirabilis, B. mufumontana, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. rubrimera, B. sangzhiensis, B. sanmingensis, B. shimentaina, B. shuichengensis, B. spinata, B. xiangnanensis, B. xianjuensis, and B. yangmingensis (vs. presence of lateral fringes on toes in these species); and from B. wushanensis (vs. presence of wide lateral fringes on toes in males while lacking in females). Having rudimentary webbing on toes, Boulenophrys yingdeensis sp. nov. differs from B. baishanzuensis, B. baolongensis, B. daweimontis, B. fansipanensis, B. frigida, B. huangshanensis, B. kuatunensis, B. lishuiensis, B. ombrophila, B. rubrimera, B. tongboensis, and B. wuliangshanensis (vs. absence of webbing on toes in these species); and from B. jingdongensis, B. palpebralespinosa, B. qianbeiensis, B. shuichengensis, and B. spinata (vs. presence of well-developed webbing on toes in these species). Description of holotype. Adult male. small body size, SVL 35.3 mm; head width slightly larger than head length, HWD/HDL 1.07; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eyes moderate in size, ED 0.30 of HDL, pupil vertical, near diamond-shaped; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanum boundary clear, ED/TD 1.68; large ovoid choanae at the base of the maxilla; vomerine ridge prominent, vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched behind; presence of a single subgular vocal sac, a pair of slit-like openings at posterior of jaw. Radio-ulnar length 0.20 of SVL and hand 0.26 of SVL; hand without webbing, fingers without lateral fringes, relative finger length I Skin of dorsum bearing small granules; skin of flanks smooth with small conical tubercles; a small, horn-like prominent tubercle on the edge of upper eyelid; clear supratympanic fold curving posteroventrally from posterior corner of eye to a level above insertion of arm; a discontinuous “X” shaped ridge on the mid-dorsum and two discontinuous dorsolateral ridges on two sides at the mid-back; sparse tubercles on the dorsal shank and thigh; ventral surface smooth; several tubercles on posterior hindlimbs; small pectoral gland closer to axilla; a single femoral positioned subequally distant from knees and cloaca on posterior surface of each thigh. Coloration of holotype in life. Dorsal surface of body olive-brown with an inverted brown triangular marking between eyes; an “X” shaped marking on the mid-dorsum. Forearms and hindlimbs with brown transverse bands. Supratympanic fold with a continuous white line; a dark vertical band below the eye, from the inferior margin of the eye to the upper lip. Ventral surface of throat and chest grayish brown with brown patches, an indistinct longitudinal stripe on surface of throat; a pair of dark brown longitudinal stripes scattered on surface of lateroventral flanks; ventral surface of body white with brown patches; ventral surface of limbs grayish brown with dark brown spots and blotches; ventral surfaces of hands and ventral surfaces of feet brown, tips of digits pale-brown; metacarpal tubercle and metatarsal tubercle reddish. Pectoral gland and femoral gland white. Iris yellowish brown. Coloration of holotype in preservative. Olive-brown faded to greyish brown dorsally. Triangular marking between eyes, “X” shaped marking on the mid-dorsum, transverse bands on dorsal forearms and hind limbs became indistinct. Color of ventral surface faded to greyish white all bands and spots became indistinct. Variation and sexual dimorphism. Measurement data of type series are listed in Table 5. Females (SVL 36.3–45.8 mm) are larger than males (SVL 33.2–35.3 mm). Presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males. Body coloration is quite variable, light brown, reddish brown, olive brown or dark brown dorsally (Figure 5). Distribution and ecology. Currently, Boulenophrys yingdeensis sp. nov. is known only from Shimentai Nature Reserve, Guangdong, China, and sympatric with B. shimentaina. All individuals were found in evergreen secondary forest, near lowland streams and nearbyleaf litter at elevations between 300– 400 m. Males perch and call on plant leaves from April to June, suggesting their breeding season corresponds to this period. Females were found on the forest floor and tadpoles were not observed.

II. Genus Pelobatrachus Beddard, 1908 Suggested common name: Clay horned toads (in English) / ���fflDzfl (in Chinese) Seven species: Pelobatrachus nasuta (Schlegel, 1858) (type species); Pelobatrachus baluensis (Boulenger, 1899); Pelobatrachus edwardinae (Inger, 1989); Pelobatrachus kalimantanensis (Munir, Hamidy, Matsui, Iskandar, Sidik & Shimada, 2019); Pelobatrachus kobayashii (Malkmus & Matsui, 1997); Pelobatrachus ligayae (Taylor, 1920); Pelobatrachus stejnegeri (Taylor, 1920)., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

IV. Genus Brachytarsophrys Tian & Hu, 1983 Suggested common name: Short-legged toads (in English) / ���DZDzfl (in Chinese), Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

3) Xenophrys lekaguli group Three species: Xenophrys lekaguli (Stuart, Chuaynkern, Chanard & Inger, 2006); Xenophrys parva (Boulenger, 1893); Xenophrys takensis (Mahony, 2011)., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979

Boulenophrys yaoshanensis sp. nov. Qi, Mo, Lyu, Wang & Wang Mt. Dayao Horned Toad / yao shan jiao chan (������fflDz) Figures 3, 7A Chresonymy. Megophrys minor ��� Liu & Hu 1962 Megophrys brachykolos ��� Fei et al. 2009; Mo et al. 2014 Boulenophrys brachykolos ��� Fei & Ye 2016; Fei 2020 Megophrys sp 31 (SYS a002189���2190, 4850���4851, 4878)��� Liu et al. 2018 Holotype. SYS a002189 (Figures 3A, 7A), adult male, collected on 8 July 2013 by Yu-Long Li and Ying-Yong Wang from the Dayaoshan Nature Reserve (26.5517��N, 114.1548��E; ca 845 m a.s.l.), Jinxiu Yao Autonomous County, Guangxi Zhuang Autonomous Region, China. Paratypes. Five adult specimens from the same locality as the holotype: male SYS a000838 collected on 13 September 2011 by Yu-Long Li and Ying-Yong Wang; males SYS a004850 and SYS a004851 / CIB 116086 collected on 1 June 2016 by Jian Wang; female SYS a004878 (Figure 3B) collected on 3 June 2016 by Jian Wang; male SYS a007023 collected on 1 June 2019 by Zhi-Tong Lyu and Yu-Long Li. Female NHMG1503016 collected on 18 March 2015 by Yun-Ming Mo from the Mt. Xianglu (24.1055��N, 110.2300��E; ca 1305 m a.s.l.), Jinxiu Yao Autonomous County; male NHMG201705032 collected on 5 May 2017 by Yun-Ming Mo from 16 km west of Jinxiu Yao Autonomous County (24.1162��N, 110.2491 ��E, ca 1115m a.s.l.), Guangxi Zhuang Autonomous Region, China. Etymology. The specific epithet yaoshanensis refers to the type locality, the Dayaoshan Nature Reserve, Guangxi, China. Particularly, we employ the epithet ���yaoshan��� rather than ���dayaoshan��� to make it consistent with other zoological and botanical species discovered from this area, e.g. Zhangixalus yaoshanensis (Liu & Hu, 1962), Leptobrachium liui yaoshanensis (Liu & Hu, 1978), Litsea yaoshanensis Yang & Huang, 1978, and Rhododendron yaoshanicum Fang & He, 1983 (Liu & Hu, 1962; Liu et al. 1978; Yang et al. 1978; Fang & He, 1983). Diagnosis. (1) Small body size, SVL 32.5���42.6 mm (37.1 �� 3.5, N = 6) in adult males and SVL 46.6���47.4 mm (47.0 �� 0.6, N = 2 in adult females; (2) snout rounded in dorsal view; (3) tympanum boundary clear, ED/TD 1.38���1.85 in males, 1.50���1.77 in females; (4) weak vomerine ridge present, vomerine teeth absent; (5) margin of tongue rounded, not notched posteriorly; (6) hind limbs slender, heels slightly overlapping or meeting, and tibiotarsal articulation reaching eye when leg stretched forward; (7) tibia 0.45���0.51 of SVL and foot 0.63���0.73 of SVL in males, tibia 0.44���0.47 of SVL and foot 0.63���0.67 of SVL in females; (8) toes without lateral fringes and with rudimentary webbing; (9) small horn-like tubercle at the edge of upper eyelid present; (10) skin of dorsum relatively smooth, a discontinuous ���X���-shaped ridge on at mid-dorsum; (11) a few sparsely distributed large tubercles on flanks; (12) body yellowish brown dorsally, an inverted hollow dark-brown triangularly shaped marking between eyes, a dark ���X���-shaped making at mid-dorsum; (13) single subgular vocal sac in males; (14) presence of villiform black nuptial spines on dorsal surface of first and second fingers in adult males. Comparisons. Comparative data of Boulenophrys yaoshanensis sp. nov. to other recognized members of the genus Boulenophrys are listed in Table 3. Having a smaller body size with SVL 32.5���42.6 mm in males, Boulenophrys yaoshanensis sp. nov. differs from the seven congeners whose SVL> 50 mm in males, including B. caudoprocta (81.3 mm in a single male), B. jingdongensis (53.0��� 56.5 mm), B. liboensis (60.5���67.7 mm), B. mirabilis (55.8���61.4 mm), B. omeimontis (56.0��� 59.5 mm), B. sangzhiensis (54.7 mm in a single male), and B. shuichengensis (102.0��� 118.3 mm). Having relatively longer shanks with heels that overlap or meet when the flexed hind limbs are held at right angles to the body axis, Boulenophrys yaoshanensis sp. nov. can be easily distinguished from the following nine congeners, B. acuta, B. brachykolos, B. daoji, B. dongguanensis, B. insularis, B. nankunensis, B. obesa, B. ombrophila, and B. wugongensis (vs. all of which have relatively shorter shanks with the heels not meeting). Lacking vomerine teeth, Boulenophrys yaoshanensis sp. nov. differs from B. caudoprocta, B. daiyunensis, B. daweimontis, B. dongguanensis, B. fansipanensis, B. frigida, B. hoanglienensis, B. insularis, B. jingdongensis, B. jinggangensis, B. jiulianensis, B. liboensis, B. nankunensis, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. rubrimera, B. sangzhiensis, B. shimentaina, and B. tongboensis (vs. presence of vomerine teeth in these species). Having an unnotched tongue, Boulenophrys yaoshanensis sp. nov. can be distinguished from B. baolongensis, B. binlingensis, B. boettgeri, B. cheni, B. hoanglienensis, B. huangshanensis, B. insularis, B. jingdongensis, B. jiulianensis, B. kuatunensis, B. liboensis, B. lushuiensis, B. minor, B. nanlingensis, B. ombrophila, B. qianbeiensis, B. sangzhiensis, B. sanmingensis, B. shuichengensis B. spinata, and B. tongboensis (vs. tongue notched posteriorly in these species). Lacking lateral fringes on toes, Boulenophrys yaoshanensis sp. nov. differs from B. acuta, B. anlongensis, B. baishanzuensis, B. binchuanensis, B. boettgeri, B. congjiangensis, B. cheni, B. daiyunensis, B. daoji, B. jingdongensis, B. jinggangensis, B. liboensis, B. lini, B. lushuiensis, B. mirabilis, B. mufumontana, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. rubrimera, B. sangzhiensis, B. sanmingensis, B. shimentaina, B. shuichengensis, B. spinata, B. xiangnanensis, B. xianjuensis, and B. yangmingensis (vs. presence of lateral fringes on toes in these species); and from B. wushanensis (vs. presence of wide lateral fringes on toes in males while lacking in females). Having rudimentary webbing on toes, Boulenophrys yaoshanensis sp. nov. differs from B. baishanzuensis, B. baolongensis, B. daweimontis, B. fansipanensis, B. frigida, B. huangshanensis, B. kuatunensis, B. lishuiensis, B. ombrophila, B. rubrimera, B. tongboensis, and B. wuliangshanensis (vs. absence of webbing on toes in these species); and from B. jingdongensis, B. palpebralespinosa, B. qianbeiensis, B. shuichengensis, and B. spinata (vs. presence of well-developed webbing on toes in these species). For the remaining seven species, Boulenophrys yaoshanensis sp. nov. can be further distinguished by the body size with SVL 46.6���47.4 mm in females (vs. 37.5���39.2 mm in B. angka, 39.5���40.4 mm in B. jiangi, 42.3 mm in B. leishanensis, 37.6 mm in B. shunhuangensis, and 50.5 mm in B. tuberogranulatus), the relative finger lengths II B. angka, IV B. caobangensis), tibio-tarsal articulation reaching eye when leg stretched forward (vs. between the nasal and tip of snout in B. shunhuangensis; between tympanum and eye in B. chishuiensis), dorsal skin relatively smooth (vs. dorsal skin rough with numerous granules in B. chishuiensis and B. tuberogranulatus), and absence of any spines on head (vs. presence of small black spines on temporal region in B. caobangensis). Description of holotype. Adult male. small body size, SVL 37.3 mm; head length equal to head width, HDW/ HDL 1.00; snout rounded in dorsal view, projecting well beyond the margin of lower jaw, sloping backward to mouth lateral in profile; top of head flat; eyes moderate in size, ED 0.38 of HDL, pupil vertical, near diamondshaped; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanum clear, ED/TD 1.68; large ovoid choanae at the base of the maxilla; weak vomerine ridge present, vomerine teeth absent, maxillary teeth present; margin of tongue rounded, not notched posteriorly; a single subgular vocal sac present, and a pair of slit-like openings at posterior of jaw. Radio-ulnar length 0.23 of SVL and hand 0.27 of SVL; hand without webbing, fingers without lateral fringes, relative finger length II Coloration of holotype in life. Dorsal surface of body yellowish brown with an inverted hollow dark-brown triangularly shaped marking between eyes; an ���X��� shaped marking on the mid-dorsum. Forearms and hind limbs with dark-brown transverse bands. Supratympanic fold light-colored; a dark vertical band below the eye, from the inferior margin of the eye to the upper lip. Ventral surface of throat and chest greyish brown with dark-brown patches; ventral surface of body greyish white with creamy white and orange spots, ventral surface of limbs greyish brown with dark brown spots; ventral surfaces of hands and feet brown, tips of digits pale brown; metacarpal tubercle and metatarsal tubercle reddish. Pectoral gland and femoral gland white. Iris yellowish brown. Coloration of holotype in preservative. Yellowish brown faded to greyish brown dorsally. Triangular marking between eyes, ���X��� shaped marking on the mid-dorsum, transverse bands on dorsal forearms and hind limbs became indistinct. Color of ventral surface faded to greyish white, all bands and spots became indistinct. Variation and sexual dimorphism. Mensural data of the type series are listed in Table 4. Two females (SYS a004878, SVL 46.6 mm; NHMG1503016, SVL 47.4 mm) are distinctly larger than the males (SVL 32.5���42.6 mm). Three paratypes (SYS a004850���4851, 4878) have subarticular tubercles on the first phalangeal articulations of the fingers. The holotype and two male paratypes (SYS a004850���4851) have small, spinose tubercles around the cloaca. Villiform black nuptial spines occur on the dorsal surface of the first and second fingers in adult males. Distribution and ecology. Currently, Boulenophrys yaoshanensis sp. nov. is known only from Jinxiu and Mengshan counties, Guangxi, China. All individuals were found in evergreen secondary forest, inhabiting flowing montane streams and the nearby forest floor and leaf litter at elevations between 800���1350 m. Males call while perched on leaves from May to July, suggesting their breeding season corresponds to this period. Females were found on the forest floor and tadpoles were not observed., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on pages 412-416, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979, {"references":["Liu, C. C. & Hu, S. C. (1962) A herpetological report of Kwangsi. Acta Zoologica Sinica, 14 (Supplement), 73 - 104. [in Chinese with English abstract]","Fei, L., Hu, S. Q., Ye, C. Y. & Huang, Y. Z. (2009) Fauna Sinica. Amphibia. Vol. 2. Anura. Science Press, Beijing, 503 pp. [in Chinese]","Mo, Y. M., Wei, Z. Y. & Chen, W. C. (2014) Colored Atlas of Guangxi Amphibians. 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5) Xenophrys vegrandis group Three species: Xenophrys vegrandis (Mahony, Teeling & Biju, 2013), Xenophrys yeae (Shi, Zhang, Xie, Jiang, Liu, Ding, Luan & Wang, 2020) comb. nov., Xenophrys zhoui (Shi, Zhang, Xie, Jiang, Liu, Ding, Luan & Wang, 2020) comb. nov.., Published as part of Qi, Shuo, Lyu, Zhi-Tong, Wang, Jian, Mo, Yun-Ming, Zeng, Zhao-Chi, Zeng, Yang-Jin, Dai, Ke-Yuan, Li, Yuan-Qiu, Grismer, L. Lee & Wang, Ying-Yong, 2021, Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China, pp. 401-438 in Zootaxa 5072 (5) on page 430, DOI: 10.11646/zootaxa.5072.5.1, http://zenodo.org/record/5748979