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Abstract In this work, we describe a new species of genus Bufo, Bufo exiguus sp. nov. from Mt. Nankun, Guangdong Province, China. This new species can be distinguished from all congeners by significant divergences in the mitochondrial 16S rRNA and CO1 genes and by a combination of morphological characters: small body size, tympanum absent, parotoid glands small and olive-shaped, tarsal fold absent, dorsal body with a fine vertebral line and white nuptial spinules present on dorsal and inner surfaces of fingers I and II in males. At present, Bufo exiguus sp. nov. is only known from the slow-flowing montane streams from its type locality and its conservation status should be carefully addressed.

The hilly region in eastern Guangdong, China lacks comprehensive scientific investigations for decades, especially in terms of herpetofauna. In recent years, several highly threatened amphibians have been gradually discovered from this region. In this work, three new species of the genus Boulenophrys are described, which are endemic from only one or two known localities in eastern Guangdong. These discoveries enrich the diversity of Boulenophrys in eastern Guangdong. With the large number of threatened urodeles and anuran species occurring in this densely populated area, the unique herpetological diversity in eastern Guangdong is facing the impacts of habitat degradation and fragmentation, and conservation actions are urgently required.

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 (2022): Erratum: SHUO QI, ZHI-TONG LYU, JIAN WANG, YUN-MING MO, ZHAO-CHI ZENG, YANG-JIN ZENG, KE-YUAN DAI, YUAN-QIU LI, L. LEE GRISMER & YING-YONG WANG (2022) Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China. Zootaxa, 5072: 401–438. Zootaxa 5115 (4): 600-600, DOI: 10.11646/zootaxa.5115.4.10, URL: http://dx.doi.org/10.11646/zootaxa.5115.4.10

Boulenophrys hungtai sp. nov. Wang, Zeng, Lyu, Xiao & Wang Hung-Ta Chang���s Horned Toad (in English) / Jiē Y��ng Ji��o Ch��n (DZffiDzdz in Chinese) Figures 5���6 Holotype. SYS a007578, adult male, collected by Jian Wang, Hong-Hui Chen and Hui-Wen Xiao on 5 January 2019 from Mt. Liwangzhang (23��38'6.42"N, 115��48'51.78"E; ca. 990 m a.s.l.), Jiexi, Jieyang, Guangdong, China. Paratypes (N=12). SYS a007575/ CIB118527, SYS a007576���7577, 7579���7582, 7594���7597, adult males, collected on 5���6 January 2019 from the same stream as the holotype at elevations between 950���1000 m. SYS a008576, adult male, collected by Jian Wang, Hong-Hui Chen, and Shuo Qi on 28 February 2021 from Shuangkeng Forestry Station (23��43'56.25"N, 116��21'26.4"E; ca. 550 m a.s.l.), Jiedong, Jieyang, Guangdong, China. Etymology. The specific epithet ��� hungtai ��� is a patronym in honor of Professor Hung-Ta Chang (=Hong-Da Zhang, �����ffl, 1914���2016), an outstanding botanist who was born in Jiexi. Diagnosis. (1) Small body size, SVL 25.8���33.3 mm (28.2 �� 2.3, N = 12) in adult males; (2) snout pointed in dorsal view; (3) tympanum moderate, TD/ED 0.52���0.61; (4) tympanic region smooth without granules or tubercles; (4) vomerine ridge and vomerine teeth absent; (5) margin of tongue rounded, not notched distally; (6) hindlimbs short, heels not meeting and tibio-tarsal articulation reaching forward to the region between tympanum and posterior corner of eye; (7) a subarticular tubercle present at the base of each fingers; (8) toes without lateral fringes and webbing; (9) distinct enlarged tubercles on the surface of limbs, flanks, chest, belly and around the cloaca; (10) tips of the tubercles on posterior abdomen, ventral thighs and around the cloaca bearing tiny spines; (11) single subgular vocal sac in males; (12) nuptial pads with villiform black nuptial spines on the dorsal surface of the first and second fingers in breeding males. Comparisons. Comparative data of Boulenophrys hungtai sp. nov. and the other recognized members of Boulenophrys are listed in Table 3. Boulenophrys hungtai sp. nov. can be easily distinguished from Boulenophrys puningensis sp. nov. by having a moderately sized tympanum, TD/ED 0.52���0.61 (vs. tympanum large, TD/ED 0.68���0.71); lacking both a vomerine ridge and vomerine teeth (vs. vomerine ridge and vomerine teeth present); lacking webbing on toes (vs. having rudimentary webbing on toes). Boulenophrys hungtai sp. nov. is strongly supported as the sister taxon to B. insularis, which is stated by Wang et al. (2017a) to be an endemic species of an offshore island in Shantou, China. Though the mean p -distance in the 16S gene is only 2.2 %, B. hungtai differs by having a snout pointed in dorsal view (vs. snout obtusely rounded in dorsal view); by the absence of vomerine ridge and vomerine teeth (vs. presence of strong vomerine ridge bearing vomerine teeth); margin of tongue not notched distally (vs. tongue notched distally); absence of webbing on toes (vs. having rudiment of webbing on toes); distinct enlarged tubercles on the surface of limbs, flanks, chest, belly and around the cloaca (vs. ventral surface smooth). Boulenophrys hungtai sp. nov. forms a clade with B. ombrophila (mean p -distance 4.9 % in the 16S gene), B. obesa (mean p -distance 3.9 % in the 16S gene), and B. cheni (mean p -distance 2.5 % in the 16S gene) though there is weak support for these relationships. However, the new species differs from the latter two congeners by having no webbing on toes (vs. having rudiment of webbing on toes in B. obesa); no vomerine ridge (vs. presence of vomerine ridges in B. obesa); a pointed snout in dorsal view (vs. snout rounded in dorsal view in B. ombrophila and B. obesa); distinct enlarged tubercles on the surface of limbs, flanks, chest, belly and around the cloaca (vs. ventral surface smooth in B. ombrophila and B. obesa); margin of tongue not notched distally, the shorter shanks with heels not meeting, absence of lateral fringes on toes (vs. tongue notched, heels meeting, presence of wide lateral fringes on toes in B. cheni). With a smaller body size, SVL 25.8���33.3 mm in adult males, Boulenophrys hungtai 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 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 a single male), and B. shuichengensis (102.0��� 118.3 mm in males). In having relatively shorter shanks with heels that do not meet when the flexed hind limbs are held at right angles to the body axis, Boulenophrys hungtai sp. nov. can be easily distinguished from the following 31 congeners, i.e. B. angka, B. anlongensis, B. baishanzuensis, B. baolongensis, B. binchuanensis, B. binlingensis, B. boettgeri, B. congjiangensis, B. chishuiensis, B. jiangi, B. jinggangensis, B. jiulianensis, B. leishanensis, B. lini, B. minor, B. mufumontana, B. nanlingensis, B. palpebralespinosa, B. qianbeiensis, B. sanmingensis, B. shimentaina, B. shunhuangensis, B. spinata, B. tongboensis, B. wuliangshanensis, B. wushanensis, B. xiangnanensis, B. xianjuensis, B. yaoshanensis, B. yangmingensis, B. yingdeensis and B. yunkaiensis, all of which have relatively longer shanks with the heels meeting or overlapping. By the absence of vomerine teeth, Boulenophrys hungtai sp. nov. differs from B. daiyunensis, B. daweimontis, B. dongguanensis, B. fansipanensis, B. frigida, B. hoanglienensis, B. nankunensis, and B. rubrimera, all of which have vomerine teeth. By having a rounded margin of the tongue that is not notched distally, Boulenophrys hungtai sp. nov. differs from B. huangshanensis, B. kuatunensis, and B. lushuiensis, all of which have notched tongues. By the absence of lateral fringes on toes, Boulenophrys hungtai sp. nov. differs from B. acuta, and B. daoji, all of which have lateral fringes on toes. By the absence of webbing on toes, Boulenophrys hungtai sp. nov. differs from B. brachykolos, B. caobangensis, B. tuberogranulatus, and B. wugongensis, all of which have rudimentary webbing on toes. Boulenophrys hungtai sp. nov. further differs from the remaining B. lishuiensis by having raised tubercles bearing spines on their tips on surface of posterior abdomen (vs. surface of belly smooth in B. lishuiensis). Description of holotype. Adult male. Body size small, SVL 28.7 mm. Head width slightly larger than head length, HDW/HDL 1.11; snout pointed 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 0.34 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 moderate in size with an obvious margin, TD/ED 0.58; large ovoid choanae at base of maxilla; vomerine ridges and vomerine teeth absent, maxillary teeth present; margin of tongue rounded, not notched distally; presence of a single subgular vocal sac, and pair of slit-like openings at posterior of jaw. Forearm (i.e., radioulna) length 0.22 of SVL and hand 0.23 of SVL; hand lacking webbing, fingers lacking lateral fringes, relative finger length I Coloration of holotype in life. Dorsal surface of body maroon, with an incomplete dark brown triangular marking between eyes. Flanks yellowish brown. Two wide oblique black bands present on forearm. Dorsal surface of fingers and hindlimbs with dark grey transverse bands. Presence of vertical dark brown band below eye. Tip of tubercle on the edge of upper eyelid white. Supratympanic fold greyish white with orange mottling. Ventral surface of throat, chest, and sides of belly dark brown with white and orange mottling, black longitudinal band on surface of throat; central and posterior part of belly white, with dark brown patches and orange mottling. Tubercles on ventral surface of chest, belly, and thighs greyish white; spines on tips of tubercles on surface of posterior abdomen, ventral surface of thighs and around cloaca dark grey. Digits, inner and outer metacarpal tubercles, and inner metatarsal tubercle greyish white. Pectoral glands and femoral glands beige. Iris reddish brown. Coloration of holotype in preservative. Maroon fades to dark brown dorsally. Coloration of flanks fades to greyish brown. Triangular marking between eyes and transverse bands on dorsal forearms and hind limbs become indistinct. Orange mottling on the supratympanic fold absent. Color of ventral surface fades; patterns of become indistinct; orange mottling on ventral skin absent. Pectoral glands and femoral glands greyish white. Variation. Mensural data of the type series are listed in Table 5. Most of the paratypes are similar to the holotype in morphology and color pattern, except for the following: coloration of dorsum is light brown and iris is reddish brown in the holotype (vs. dorsum beige and iris greyish white with irregular dark brown and light orange patterns in the paratype SYS a008576 (Fig. 6A���B); central and posterior part of belly white, with dark brown patches and orange mottling in the holotype (vs. ventral skin dark brown, without regular patches in the paratype SYS a008576 (Fig. 6C���D); larger body size in the paratypes SYS a007582 (SVL 32.7 mm) and SYS 008576 (SVL 33.3 mm); tibio-tarsal articulation reaching forward to middle of tympanum when hind limb stretched along body (vs. tibio-tarsal articulation reaching forward to posterior corner of eye in the paratypes SYS a007576, 7582, 7596, 7597, 8576). Distribution and natural history. Currently, Boulenophrys hungtai sp. nov. is known from Mt. Liwangzhang of Jiexi (800���1000 m a.s.l.) and Shuangkeng Forestry Station (500���800 m a.s.l.) of Jiedong, which are ca. 55 km from each other. This toad inhabits flowing montane streams and the nearby forest floor and leaf litter, and is sympatric with Pachytriton brevipes (Sauvage, 1876). Cynops orphicus Risch, 1983 can also be observed in the surrounding area. Advertisement calls of males from Mt. Liwangzhang were heard from November to the following January. A single male (SYS a008576) from Shuangkeng Forestry Station was noticed discontinuously calling on March after the heavy rain. Males were found calling under the leaf litter or in rock crevices in flowing streams., Published as part of Wang, Jian, Zeng, Zhao-Chi, Lyu, Zhi-Tong, Qi, Shuo, Liu, Zu-Yao, Chen, Hong-Hui, Lu, Yu-Hong, Xiao, Hui-Wen, Lin, Can-Rong, Chen, Kai & Wang, Ying-Yong, 2022, Description of three new Boulenophrys species from eastern Guangdong, China, emphasizing the urgency of ecological conservation in this region (Anura, Megophryidae), pp. 91-119 in Zootaxa 5099 (1) on pages 106-110, DOI: 10.11646/zootaxa.5099.1.4, http://zenodo.org/record/6036964, {"references":["Wang, J., Liu, Z. Y., Lyu, Z. T., Zeng, Z. C. & Wang, Y. Y. (2017 a) A new species of the genus Xenophrys (Amphibia: Anura: Megophryidae) from an offshore island in Guangdong Province, southeastern China. Zootaxa, 4324 (3), 541 - 556. https: // doi. org / 10.11646 / zootaxa. 4324.3.8","Sauvage, H. E. (1876) L'Institut. Journal des Academies et Societes Scientifiques de la France et de l'Etrangers, Paris, 4, 274 - 275.","Risch, J. P. (1983) Cynops orphicus, a new salamander from Guangdong Province, South China (Amphibia, Caudata, Salamandridae). Alytes, Paris, 2, 45 - 52."]}

Boulenophrys fengshunensis sp. nov. Wang, Zeng, Lyu & Wang Fengshun Horned Toad (in English) / Fēng Sh��n Ji��o Ch��n (���NJDzdz in Chinese) Chresonymy: Megophrys sp 14 ��� Liu et al. 2018 Figures 7���8 Holotype. SYS a004744, adult male, collected by Jian Wang and Zhi-Tong Lyu on 13 May 2016 from Mt. Tongguzhang (24��10'31.12"N, 116��21'2.63"E; ca. 1500 m a.s.l.), Fengshun, Meizhou, Guangdong, China. Paratypes (N=7). Adult males, SYS a004724/ CIB118528, SYS a004725���4728, collected by Jian Wang and Zhi-Tong Lyu on 11 May 2016 from the same locality of the holotype. Adult females, SYS a005220���5221, collected by Jian Wang on 9 August 2016 from the same locality of the holotype. Etymology. The specific epithet ��� fengshunensis ��� refers to the type locality of the new species, the Fengshun County. Diagnosis. (1) Small body size, SVL 34.3���39.4 mm (36.9 �� 1.7, N = 6) in adult males and SVL 42.5���44.9 mm (N = 2) in adult females; (2) snout pointed in dorsal view; (3) tympanum moderate in size TD/ED 0.56���0.67, tubercles bearing spines on tips present on skin of temporal region including the tympanum; (4) vomerine ridge and vomerine teeth present; (5) margin of tongue rounded, not notched distally; (6) hindlimbs short, heels not meeting and tibio-tarsal articulation reaching forward to the region between tympanum and posterior corner of eye, TIB/SVL 0.37���0.43; (7) a subarticular tubercle present at the base of each fingers; (8) toes without lateral fringes and with rudiment of webbing; (9) distinct sparse enlarged tubercles on the surface of limbs, flanks, posterior part of belly and around the cloaca; (10) tips of the tubercles on ventral surface of thighs and around the cloaca bearing tiny spines; (11) single subgular vocal sac in males; (12) nuptial pads with well-developed villiform black nuptial spines on the dorsal surface of the first and second fingers in breeding males. Comparisons. Comparative data of Boulenophrys fengshunensis sp. nov. and the other recognized members of Boulenophrys are listed in Table 3. Boulenophrys fengshunensis sp. nov. can be distinguished from Boulenophrys puningensis sp. nov. and Boulenophrys hungtai sp. nov. having a tympanum with tubercles bearing spines on their tips (vs. tympanic region smooth without granules, tubercles or spines in Boulenophrys puningensis sp. nov. and Boulenophrys hungtai sp. nov.); the presence of both a vomerine ridge and vomerine teeth (vs. absence of vomerine ridge but with vomerine teeth present in Boulenophrys hungtai sp. nov.); tubercles on skin of the posterior part of belly without spines (vs. tubercles on skin of the posterior part of belly bearing spines on their tips in both Boulenophrys puningensis sp. nov. and Boulenophrys hungtai sp. nov.). With a smaller body size, SVL 34.3���39.4 mm in adult males and SVL 42.5���44.9 mm in adult females, Boulenophrys fengshunensis sp. nov. differs from the seven congeners whose SVL ��� 50 mm in males or SVL ��� 60 mm in females, including B. caudoprocta (81.3 mm in a single male), B. jingdongensis (53.0��� 56.5 mm in males and 63.5 mm in a single female), B. liboensis (60.5���67.7 mm in males and 60.8���70.6 in females), B. mirabilis (55.8���61.4 mm in males and 68.5���74.8 mm in females), B. omeimontis (56.0��� 59.5 mm in males and 68.0��� 72.5 mm in females), B. sangzhiensis (54.7 mm in a single male), and B. shuichengensis (102.0��� 118.3 mm in males and 99.8���115.6 mm in females). In having relatively shorter shanks with heels that do not meet when the flexed hind limbs are held at right angles to the body axis, Boulenophrys fengshunensis sp. nov. can be easily distinguished from the following 35 congeners, i.e. B. angka, B. anlongensis, B. baishanzuensis, B. baolongensis, B. binchuanensis, B. binlingensis, B. boettgeri, B. congjiangensis, B. cheni, B. chishuiensis, B. daiyunensis, B. jiangi, B. jinggangensis, B. jiulianensis, B. leishanensis, B. lini, B. minor, B. mufumontana, B. nanlingensis, B. palpebralespinosa, B. qianbeiensis, B. sanmingensis, B. shimentaina, B. shunhuangensis, B. spinata, B. tongboensis, B. tuberogranulatus, B. wuliangshanensis, B. wushanensis, B. xiangnanensis, B. xianjuensis, B. yaoshanensis, B.yangmingensis, B. yingdeensis and B. yunkaiensis, all of which have relatively longer shanks with the heels meeting or overlapping. By the presence of vomerine teeth, Boulenophrys fengshunensis sp. nov. differs from B. acuta, B. brachykolos, B. caobangensis, B. daoji, B. huangshanensis, B. kuatunensis, B. lishuiensis, B. lushuiensis, B. obesa, B. ombrophila, and B. wugongensis, all of which lack vomerine teeth. By having a rounded margin of the tongue that is not notched distally, Boulenophrys fengshunensis sp. nov. differs from B. hoanglienensis, B. insularis, which have a notched tongue. By the absence of lateral fringes on toes, Boulenophrys fengshunensis sp. nov. differs from B. rubrimera, which has narrow lateral fringes on toes. By the presence of rudimentary webbing on toes, Boulenophrys fengshunensis sp. nov. differs from B. daweimontis, B. fansipanensis, B. frigida, and B. rubrimera, all of which lack webbing on toes. Boulenophrys fengshunensi sp. nov. further differs from the remaining B. dongguanensis and B. nankunensis by having raised tubercles on the surface of posterior abdomen (vs. absence of such tubercles in both B. dongguanensis B. nankunensis); the presence of tubercles bearing spines on their tips on the temporal region including the tympanum (vs. temporal region lacking tubercles or spines in both B. dongguanensis and B. nankunensis). Description of holotype. Adult male. Small body size, SVL 37.4 mm; head width slightly larger than head length, HWD/HDL 1.17; snout pointed 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 0.37 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 moderate in size with an obvious margin, TD/ED 0.65; large ovoid choanae at base of maxilla; vomerine ridge and vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched distally; presence of single subgular vocal sac, and pair of slit-like openings at posterior of jaw. Forearm (i.e., radioulna) length 0.21 of SVL and hand 0.25 of SVL; hand without webbing, fingers without lateral fringes, relative finger length I = II Coloration of holotype in life. Dorsal surface of body dark brown, with incomplete dark brown triangular marking between eyes. Two wide oblique black bands present on forearm. Dorsal surface of fingers and hindlimbs with dark grey transverse bands. Presence of vertical dark brown band below eye. Tip of tubercles on edge of upper eyelid greyish white. Supratympanic fold orangey brown. Ventral surface dark brown, black longitudinal band on surface of throat, posterior part of belly with irregular greyish white patches. Tubercles on ventral surface of posterior part of belly and thighs greyish white; spines on tips of tubercles on temporal region, ventral surface of thighs and around cloaca dark grey. Digits, inner and outer metacarpal tubercles, and inner metatarsal tubercle dark grey. Pectoral glands beige and femoral glands greyish white. Iris orangish brown. Coloration of holotype in preservative. Dorsal surface of body dark brown. Triangular marking between eyes, vertical dark brown band below eye and transverse bands on dorsal forearms and hind limbs become indistinct. Supratympanic fold greyish white. All bands and patterns on ventral surface no longer apparent. Irregular greyish white patches on posterior part of belly become dark grey. Variation. Mensural data of the type series are listed in Table 6. Most of the paratypes are similar to the holotype in morphology and color pattern, except for the following: dorsal surface and ventral surface dark brown, posterior part of belly with irregular greyish white patches, ventral surface of thighs dark grey, triangular marking between eyes is incomplete in the holotype (vs. dorsal surface beige and ventral surface light orange, posterior part of belly and ventral surface of thighs greyish white, triangular marking between eyes is complete in the female paratype SYS a005221 (Fig. 8); tibio-tarsal articulation reaching forward to middle of tympanum when hind limb stretched along body in the holotype (vs. tibio-tarsal articulation reaching forward to posterior margin of tympanum in the paratype SYS a004725; reaching forward to the region between tympanum and eye in the paratype SYS a004726). Females are distinctly larger than the males. Distribution and natural history. Currently, Boulenophrys fengshunensis sp. nov. is only known from its type locality, Mt. Tongguzhang of Fengshun. It inhabits flowing montane streams and the nearby forest floor and leaf litter at elevations between 800���1500 m. Boulenophrys fengshunensis sp. nov. is found to be sympatric with Pachytriton granulosus Chang, 1933 and Cynops glaucus Yuan, Jiang, Ding, Zhang & Che, 2013. Advertisement calls of males were heard during April and June. Males were found calling under the leaf litters around the flowing seeps. Tadpoles could be found in this period. Female specimens collected during August had immature eggs; males were not heard calling during this period, but sub-adults were observed., Published as part of Wang, Jian, Zeng, Zhao-Chi, Lyu, Zhi-Tong, Qi, Shuo, Liu, Zu-Yao, Chen, Hong-Hui, Lu, Yu-Hong, Xiao, Hui-Wen, Lin, Can-Rong, Chen, Kai & Wang, Ying-Yong, 2022, Description of three new Boulenophrys species from eastern Guangdong, China, emphasizing the urgency of ecological conservation in this region (Anura, Megophryidae), pp. 91-119 in Zootaxa 5099 (1) on pages 110-114, DOI: 10.11646/zootaxa.5099.1.4, http://zenodo.org/record/6036964, {"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","Chang, M. L. Y. (1933) On the salamanders of Chekiang. Contributions from the Biological Laboratory of the Science Society of China. Zoological Series, 9, 305 - 328.","Yuan, Z. Y., Jiang, K., Ding, L., Zhang, L. & Che, J. (2013) A new newt of the genus Cynops (Caudata: Salamandridae) from Guangdong, China. Asian Herpetological Research, 4 (2), 116 - 123. https: // doi. org / 10.3724 / SP. J. 1245.2013.00116"]}

Boulenophrys puningensis sp. nov. Wang, Zeng, Lyu, Xiao & Wang Puning Horned Toad (in English) / P�� N��ng Ji��o Ch��n (D���Dzdz in Chinese) Figures 3���4 Holotype. SYS a005770, adult male, collected by Jian Wang on 24April 2017 from Longkeng Village (23��7'54.07"N, 115��51'5.28"E; ca. 120 m a.s.l.), Daping Town, Puning, Jieyang, Guangdong, China. Paratypes (N=5). Adult male, SYS a006755/ CIB118526, collected by Jian Wang, Can-Rong Lin and Hui-Wen Xiao on 14 February 2018; adult males SYS a007649, 7650 and adult females SYS a007647, 7648, collected by Jian Wang, Can-Rong Lin and Hui-Wen Xiao on 18 March 2019, all from the same stream as the holotype at elevations between 250���300 m. Etymology. The specific epithet ��� puningensis ��� refers to the type locality of the new species in Puning. Three of the authors of this work (Jian Wang, Hui-Wen Xiao and Can-Rong Lin) chose this nomen in honor of their hometown. Diagnosis. (1) Small body size, SVL 31.7���34.6 mm (33.0 �� 1.3, N = 4) in adult males and SVL 37.8���38.3 mm (N = 2) in adult females; (2) snout rounded in dorsal view; (3) tympanum large, TD/ED 0.68���0.71; (4) tympanic region smooth without granules or tubercles; (5) vomerine ridge and vomerine teeth present; (6) margin of tongue rounded, not notched distally; (7) hindlimbs short, heels not meeting and tibio-tarsal articulation reaching forward to the region between tympanum and eye; (8) a subarticular tubercle present at the base of each fingers; (9) toes without lateral fringes and with rudiment of webbing; (10) distinct enlarged tubercles on the surface of limbs, flanks, chest, belly and around the cloaca; (11) tips of the enlarged tubercles on posterior abdomen, ventral thighs and around the cloaca bearing tiny spines; (12) single subgular vocal sac in males; (13) nuptial pads with villiform black nuptial spines on the dorsal surface of the first and second fingers in breeding males. Comparisons. Comparative data of Boulenophrys puningensis sp. nov. and the other recognized members of Boulenophrys are listed in Table 3. With a smaller body size, SVL 31.7���34.6 mm in adult males and SVL 37.8���38.3 mm in adult females, Boulenophrys puningensis sp. nov. differs from the eight congeners whose SVL ��� 50 mm in adult males or females, including B. caudoprocta (81.3 mm in a single male), B. jingdongensis (53.0��� 56.5 mm in males and 63.5 mm in a single female), B. liboensis (60.5���67.7 mm in males and 60.8���70.6 in females), B. mirabilis (55.8���61.4 mm in males and 68.5���74.8 mm in females), B. omeimontis (56.0��� 59.5 mm in males and 68.0��� 72.5 mm in females), B. sangzhiensis (54.7 mm in a single male), B. shuichengensis (102.0��� 118.3 mm in males and 99.8���115.6 mm in females), and B. spinata (54.0���55.0 mm in females). Boulenophrys puningensis sp. nov. shows the least genetic divergence from B. kuatunensis (mean p -distances 5.3 % in the 16S gene) and B. daiyunensis (mean p -distances 6.2 % in the 16S gene). However, the new species distinctively differs from these species by having relatively shorter shanks with the heels not meeting when the flexed hindlimbs are held at right angles to the body axis (vs. heels meeting or overlapping in B. daiyunensis); having rudiment of webbing and no lateral fringes on toes (vs. lateral fringes narrow in B. daiyunensis; webbing absent in B. kuatunensis); having raised and enlarged tubercles with spines on their tips on surface of posterior abdomen, ventral thighs and around the cloaca (vs. such tubercles not enlarged and without spines in B. daiyunensis; ventral surface smooth in B. kuatunensis). Boulenophrys puningensis sp. nov. is morphologically most similar to B. brachykolos, which is restricted to Hong Kong and Shenzhen, China (Liu et al. 2018). The new species differs from B. brachykolos by having vomerine teeth (vs. absent in B. brachykolos); lacking spines on the surface of the tympanic region (vs. having dense tiny spines on the surface of the tympanic region in B. brachykolos); and having different relative finger length formula (I = II Banophrys puningensis sp. nov. vs. II B. brachykolos). In having relatively shorter shanks with heels that do not meet when the flexed hind limbs are held at right angles to the body axis, Boulenophrys puningensis sp. nov. can be easily distinguished from the following 32 congeners, i.e. B. angka, B. anlongensis, B. baishanzuensis, B. baolongensis, B. binchuanensis, B. binlingensis, B. boettgeri, B. congjiangensis, B. cheni, B. chishuiensis, B. jiangi, B. jinggangensis, B. jiulianensis, B. leishanensis, B. lini, B. minor, B. mufumontana, B. nanlingensis, B. palpebralespinosa, B. qianbeiensis, B. sanmingensis, B. shimentaina, B. shunhuangensis, B. tongboensis, B. wuliangshanensis, B. wushanensis, B. xiangnanensis, B. xianjuensis, B. yaoshanensis, B. yangmingensis, B. yingdeensis and B. yunkaiensis, all of which have relatively longer shanks with the heels meeting or overlapping. By the presence of vomerine teeth, Boulenophrys puningensis sp. nov. differs from B. acuta, B. caobangensis, B. daoji, B. huangshanensis, B. lishuiensis, B. lushuiensis, B. obesa, B. ombrophila, B. tuberogranulatus, and B. wugongensis, all of which lack vomerine teeth. By having a rounded tongue margin that is not notched distally, Boulenophrys puningensis sp. nov. differs from B. hoanglienensis, and B. insularis, all of which have notched tongues. By the absence of lateral fringes on toes, Boulenophrys puningensis sp. nov. differs from B. rubrimera, which has narrow lateral fringes on toes. By the presence of rudimentary webbing on the toes, Boulenophrys puningensis sp. nov. differs from B. daweimontis, B. fansipanensis, and B. frigida, all of which lack webbing on the toes. Boulenophrys puningensis sp. nov. further differs from the remaining B. dongguanensis and B. nankunensis by having raised tubercles bearing spines on the surface of the posterior abdomen, ventral thighs, and around the cloaca (vs. absence of such tubercles and spines in B. dongguanensis and B. nankunensis). Description of holotype. Adult male. Body size small, SVL 34.6 mm. Head width slightly larger than head length, HWD/HDL 1.03; 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 0.36 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 large with an obvious margin, TD/ED 0.70; large ovoid choanae at base of maxilla; vomerine ridge and vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched distally; presence of single subgular vocal sac, and pair of slit-like openings at posterior of jaw. Forearm (i.e., radioulna) length 0.21 of SVL and hand 0.23 of SVL; hand without webbing, fingers without lateral fringes, relative finger length I = II Coloration of holotype in life. Dorsal surface of body yellowish brown, with incomplete dark brown triangular marking between eyes. Two wide oblique black bands present on forearm. Dorsal surface of fingers and hindlimbs with dark grey transverse bands. Presence of vertical dark brown band below eye. Tubercles on edge of upper eyelid beige. Supratympanic fold light brown. Ventral surface dark grey, with black longitudinal band on surface of throat; surface of throat and chest mottled with orange patches. Tubercles on ventral surface of chest, belly, and thighs greyish white; spines on tips of tubercles on surface of posterior abdomen; ventral surface of thighs and around cloaca dark grey. Digits, inner and outer metacarpal tubercles and inner metatarsal tubercle greyish white. Pectoral glands and femoral glands beige, mottled with orange patches. Iris yellowish brown, with greyish white patches on upper and lower margin. Coloration of holotype in preservative. Yellowish brown fades to greyish brown dorsally. Color of the triangular marking between eyes, oblique bands on forearms, patterns on ventral surface faded. Orange patches on surface of throat, chest; color of pectoral glands and femoral glands faded. Variation. Mensural data of the type series are listed in Table 4. Most of the paratypes are similar to the holotype in morphology and color pattern, except for the following: dorsal surface of body yellowish brown in the holotype (vs. dorsal surface of body light brown in the paratypes SYS a007647 (Fig. 4C) and SYS a007648 (Fig. 4E); ventral surface dark grey with orange patches (vs. ventral surface lacking bright patches in the paratypes SYS a007649 (Fig. 4B), SYS a007647 (Fig. 4D) and SYS a007648 (Fig. 4F); iris yellowish brown, with greyish white patches on its upper and lower margin in the holotype (vs. iris grey with beige and dark mottling in the paratype SYS a007649 (Fig. 4A); tubercles on posterior part of abdomen of the paratype SYS a007648 (Fig. 4F) are weakly developed. Females are distinctly larger than the males. Distribution and natural history. Currently, Boulenophrys puningensis sp. nov. is only known from its type locality, Longkeng Village of Puning. It inhabits flowing montane streams and the nearby forest floor and leaf litter at elevations between 120��� 300 m. Advertisement calls of males were heard from February until April. Males were found calling in rock crevices in the flowing streams. Tadpoles could be found in this period., Published as part of Wang, Jian, Zeng, Zhao-Chi, Lyu, Zhi-Tong, Qi, Shuo, Liu, Zu-Yao, Chen, Hong-Hui, Lu, Yu-Hong, Xiao, Hui-Wen, Lin, Can-Rong, Chen, Kai & Wang, Ying-Yong, 2022, Description of three new Boulenophrys species from eastern Guangdong, China, emphasizing the urgency of ecological conservation in this region (Anura, Megophryidae), pp. 91-119 in Zootaxa 5099 (1) on pages 102-106, DOI: 10.11646/zootaxa.5099.1.4, http://zenodo.org/record/6036964, {"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"]}

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

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

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

VIII. Incertae sedis with Megophryinae Two species: ��� Megophrys ��� dringi Inger, Stuebing & Tan, 1995; ��� Megophrys ��� feii Yang, Wang & Wang, 2018., 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

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

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. Guangxi Science & Technology Publishing House, Nanning, 282 pp. [in Chinese]","Fei, L. & Ye, C. Y. (2016) Amphibians of China. (Vol. 1). Science Press, Beijing.","Fei, L. (2020) Atlas of Amphibians in China. Field Edition. Henan Science and Technology Press, Zhengzhou, 432 pp. [in Chinese]","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","Liu, C. C., Hu, S. Q., Tian, W. S. & Wu, G. F. (1978) Four new species of amphibians from Sichuan and Guangxi. Materials for Herpetological Research, 4, 18 - 19. [in Chinese]","Yang, Y. C., Huang, P. H., Tsui, H. P., Li, H. W. & Pai, P. Y. (1978) Materiae ad Floram Lauracearum Sinicarum (II). Acta Phytotaxonomica Sinica, 16 (4), 38 - 69. [in Chinese]","Fang, W. P. & He, M. Y. (1983) Studies on the genus Rhododendron (S). Bulletin of Botanical Research, 3 (1), 1 - 8. [in Chinese]"]}

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

Boulenophrys fengshunensis sp. nov. Wang, Zeng, Lyu & Wang Fengshun Horned Toad (in English) / Fēng Shùn Jiǎo Chán (丰NJDzdz in Chinese) Chresonymy: Megophrys sp 14 — Liu et al. 2018 Figures 7–8 Holotype. SYS a004744, adult male, collected by Jian Wang and Zhi-Tong Lyu on 13 May 2016 from Mt. Tongguzhang (24°10'31.12"N, 116°21'2.63"E; ca. 1500 m a.s.l.), Fengshun, Meizhou, Guangdong, China. Paratypes (N=7). Adult males, SYS a004724/ CIB118528, SYS a004725–4728, collected by Jian Wang and Zhi-Tong Lyu on 11 May 2016 from the same locality of the holotype. Adult females, SYS a005220–5221, collected by Jian Wang on 9 August 2016 from the same locality of the holotype. Etymology. The specific epithet “ fengshunensis ” refers to the type locality of the new species, the Fengshun County. Diagnosis. (1) Small body size, SVL 34.3–39.4 mm (36.9 ± 1.7, N = 6) in adult males and SVL 42.5–44.9 mm (N = 2) in adult females; (2) snout pointed in dorsal view; (3) tympanum moderate in size TD/ED 0.56–0.67, tubercles bearing spines on tips present on skin of temporal region including the tympanum; (4) vomerine ridge and vomerine teeth present; (5) margin of tongue rounded, not notched distally; (6) hindlimbs short, heels not meeting and tibio-tarsal articulation reaching forward to the region between tympanum and posterior corner of eye, TIB/SVL 0.37–0.43; (7) a subarticular tubercle present at the base of each fingers; (8) toes without lateral fringes and with rudiment of webbing; (9) distinct sparse enlarged tubercles on the surface of limbs, flanks, posterior part of belly and around the cloaca; (10) tips of the tubercles on ventral surface of thighs and around the cloaca bearing tiny spines; (11) single subgular vocal sac in males; (12) nuptial pads with well-developed villiform black nuptial spines on the dorsal surface of the first and second fingers in breeding males. Comparisons. Comparative data of Boulenophrys fengshunensis sp. nov. and the other recognized members of Boulenophrys are listed in Table 3. Boulenophrys fengshunensis sp. nov. can be distinguished from Boulenophrys puningensis sp. nov. and Boulenophrys hungtai sp. nov. having a tympanum with tubercles bearing spines on their tips (vs. tympanic region smooth without granules, tubercles or spines in Boulenophrys puningensis sp. nov. and Boulenophrys hungtai sp. nov.); the presence of both a vomerine ridge and vomerine teeth (vs. absence of vomerine ridge but with vomerine teeth present in Boulenophrys hungtai sp. nov.); tubercles on skin of the posterior part of belly without spines (vs. tubercles on skin of the posterior part of belly bearing spines on their tips in both Boulenophrys puningensis sp. nov. and Boulenophrys hungtai sp. nov.). With a smaller body size, SVL 34.3–39.4 mm in adult males and SVL 42.5–44.9 mm in adult females, Boulenophrys fengshunensis sp. nov. differs from the seven congeners whose SVL ≥ 50 mm in males or SVL ≥ 60 mm in females, including B. caudoprocta (81.3 mm in a single male), B. jingdongensis (53.0– 56.5 mm in males and 63.5 mm in a single female), B. liboensis (60.5–67.7 mm in males and 60.8–70.6 in females), B. mirabilis (55.8–61.4 mm in males and 68.5–74.8 mm in females), B. omeimontis (56.0– 59.5 mm in males and 68.0– 72.5 mm in females), B. sangzhiensis (54.7 mm in a single male), and B. shuichengensis (102.0– 118.3 mm in males and 99.8–115.6 mm in females). In having relatively shorter shanks with heels that do not meet when the flexed hind limbs are held at right angles to the body axis, Boulenophrys fengshunensis sp. nov. can be easily distinguished from the following 35 congeners, i.e. B. angka, B. anlongensis, B. baishanzuensis, B. baolongensis, B. binchuanensis, B. binlingensis, B. boettgeri, B. congjiangensis, B. cheni, B. chishuiensis, B. daiyunensis, B. jiangi, B. jinggangensis, B. jiulianensis, B. leishanensis, B. lini, B. minor, B. mufumontana, B. nanlingensis, B. palpebralespinosa, B. qianbeiensis, B. sanmingensis, B. shimentaina, B. shunhuangensis, B. spinata, B. tongboensis, B. tuberogranulatus, B. wuliangshanensis, B. wushanensis, B. xiangnanensis, B. xianjuensis, B. yaoshanensis, B.yangmingensis, B. yingdeensis and B. yunkaiensis, all of which have relatively longer shanks with the heels meeting or overlapping. By the presence of vomerine teeth, Boulenophrys fengshunensis sp. nov. differs from B. acuta, B. brachykolos, B. caobangensis, B. daoji, B. huangshanensis, B. kuatunensis, B. lishuiensis, B. lushuiensis, B. obesa, B. ombrophila, and B. wugongensis, all of which lack vomerine teeth. By having a rounded margin of the tongue that is not notched distally, Boulenophrys fengshunensis sp. nov. differs from B. hoanglienensis, B. insularis, which have a notched tongue. By the absence of lateral fringes on toes, Boulenophrys fengshunensis sp. nov. differs from B. rubrimera, which has narrow lateral fringes on toes. By the presence of rudimentary webbing on toes, Boulenophrys fengshunensis sp. nov. differs from B. daweimontis, B. fansipanensis, B. frigida, and B. rubrimera, all of which lack webbing on toes. Boulenophrys fengshunensi sp. nov. further differs from the remaining B. dongguanensis and B. nankunensis by having raised tubercles on the surface of posterior abdomen (vs. absence of such tubercles in both B. dongguanensis B. nankunensis); the presence of tubercles bearing spines on their tips on the temporal region including the tympanum (vs. temporal region lacking tubercles or spines in both B. dongguanensis and B. nankunensis). Description of holotype. Adult male. Small body size, SVL 37.4 mm; head width slightly larger than head length, HWD/HDL 1.17; snout pointed 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 0.37 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 moderate in size with an obvious margin, TD/ED 0.65; large ovoid choanae at base of maxilla; vomerine ridge and vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched distally; presence of single subgular vocal sac, and pair of slit-like openings at posterior of jaw. Forearm (i.e., radioulna) length 0.21 of SVL and hand 0.25 of SVL; hand without webbing, fingers without lateral fringes, relative finger length I = II Dorsal skin rough, granular, with sparse tubercles; skin of dorsal body and limbs bearing tiny spines; sparse large conical tubercles on flanks; single horn-like prominent tubercle on edge of upper eyelid; obvious supratympanic fold curving posteroventrally from posterior corner of eye to level above insertion of arm; dense tubercles on skin of upper lip, upper eyelid, mandibular articulation, loreal, temporal region including tympanum and surface around cloaca; tip of tubercles on skin of temporal region excluding tympanum bearing tiny spine; single discontinuous “X” shaped ridge and few elongated tubercles arranged in longitudinal rows on two sides at mid-dorsum; sparse tubercles on dorsal shank and thigh; ventral skin of throat, chest and anterior part of belly smooth; ventral skin of thighs and posterior part of belly bearing dense tubercles; surface around cloaca with dense tubercles bearing tiny spines; small pectoral gland closer to axilla; single femoral gland positioned on posterior surface of thigh at midpoint between knee and cloaca.. Coloration of holotype in life. Dorsal surface of body dark brown, with incomplete dark brown triangular marking between eyes. Two wide oblique black bands present on forearm. Dorsal surface of fingers and hindlimbs with dark grey transverse bands. Presence of vertical dark brown band below eye. Tip of tubercles on edge of upper eyelid greyish white. Supratympanic fold orangey brown. Ventral surface dark brown, black longitudinal band on surface of throat, posterior part of belly with irregular greyish white patches. Tubercles on ventral surface of posterior part of belly and thighs greyish white; spines on tips of tubercles on temporal region, ventral surface of thighs and around cloaca dark grey. Digits, inner and outer metacarpal tubercles, and inner metatarsal tubercle dark grey. Pectoral glands beige and femoral glands greyish white. Iris orangish brown. Coloration of holotype in preservative. Dorsal surface of body dark brown. Triangular marking between eyes, vertical dark brown band below eye and transverse bands on dorsal forearms and hind limbs become indistinct. Supratympanic fold greyish white. All bands and patterns on ventral surface no longer apparent. Irregular greyish white patches on posterior part of belly become dark grey. Variation. Mensural data of the type series are listed in Table 6. Most of the paratypes are similar to the holotype in morphology and color pattern, except for the following: dorsal surface and ventral surface dark brown, posterior part of belly with irregular greyish white patches, ventral surface of thighs dark grey, triangular marking between eyes is incomplete in the holotype (vs. dorsal surface beige and ventral surface light orange, posterior part of belly and ventral surface of thighs greyish white, triangular marking between eyes is complete in the female paratype SYS a005221 (Fig. 8); tibio-tarsal articulation reaching forward to middle of tympanum when hind limb stretched along body in the holotype (vs. tibio-tarsal articulation reaching forward to posterior margin of tympanum in the paratype SYS a004725; reaching forward to the region between tympanum and eye in the paratype SYS a004726). Females are distinctly larger than the males. Distribution and natural history. Currently, Boulenophrys fengshunensis sp. nov. is only known from its type locality, Mt. Tongguzhang of Fengshun. It inhabits flowing montane streams and the nearby forest floor and leaf litter at elevations between 800–1500 m. Boulenophrys fengshunensis sp. nov. is found to be sympatric with Pachytriton granulosus Chang, 1933 and Cynops glaucus Yuan, Jiang, Ding, Zhang & Che, 2013. Advertisement calls of males were heard during April and June. Males were found calling under the leaf litters around the flowing seeps. Tadpoles could be found in this period. Female specimens collected during August had immature eggs; males were not heard calling during this period, but sub-adults were observed.

Boulenophrys puningensis sp. nov. Wang, Zeng, Lyu, Xiao & Wang Puning Horned Toad (in English) / Pǔ Níng Jiǎo Chán (DṪDzdz in Chinese) Figures 3–4 Holotype. SYS a005770, adult male, collected by Jian Wang on 24April 2017 from Longkeng Village (23°7'54.07"N, 115°51'5.28"E; ca. 120 m a.s.l.), Daping Town, Puning, Jieyang, Guangdong, China. Paratypes (N=5). Adult male, SYS a006755/ CIB118526, collected by Jian Wang, Can-Rong Lin and Hui-Wen Xiao on 14 February 2018; adult males SYS a007649, 7650 and adult females SYS a007647, 7648, collected by Jian Wang, Can-Rong Lin and Hui-Wen Xiao on 18 March 2019, all from the same stream as the holotype at elevations between 250–300 m. Etymology. The specific epithet “ puningensis ” refers to the type locality of the new species in Puning. Three of the authors of this work (Jian Wang, Hui-Wen Xiao and Can-Rong Lin) chose this nomen in honor of their hometown. Diagnosis. (1) Small body size, SVL 31.7–34.6 mm (33.0 ± 1.3, N = 4) in adult males and SVL 37.8–38.3 mm (N = 2) in adult females; (2) snout rounded in dorsal view; (3) tympanum large, TD/ED 0.68–0.71; (4) tympanic region smooth without granules or tubercles; (5) vomerine ridge and vomerine teeth present; (6) margin of tongue rounded, not notched distally; (7) hindlimbs short, heels not meeting and tibio-tarsal articulation reaching forward to the region between tympanum and eye; (8) a subarticular tubercle present at the base of each fingers; (9) toes without lateral fringes and with rudiment of webbing; (10) distinct enlarged tubercles on the surface of limbs, flanks, chest, belly and around the cloaca; (11) tips of the enlarged tubercles on posterior abdomen, ventral thighs and around the cloaca bearing tiny spines; (12) single subgular vocal sac in males; (13) nuptial pads with villiform black nuptial spines on the dorsal surface of the first and second fingers in breeding males. Comparisons. Comparative data of Boulenophrys puningensis sp. nov. and the other recognized members of Boulenophrys are listed in Table 3. With a smaller body size, SVL 31.7–34.6 mm in adult males and SVL 37.8–38.3 mm in adult females, Boulenophrys puningensis sp. nov. differs from the eight congeners whose SVL ≥ 50 mm in adult males or females, including B. caudoprocta (81.3 mm in a single male), B. jingdongensis (53.0– 56.5 mm in males and 63.5 mm in a single female), B. liboensis (60.5–67.7 mm in males and 60.8–70.6 in females), B. mirabilis (55.8–61.4 mm in males and 68.5–74.8 mm in females), B. omeimontis (56.0– 59.5 mm in males and 68.0– 72.5 mm in females), B. sangzhiensis (54.7 mm in a single male), B. shuichengensis (102.0– 118.3 mm in males and 99.8–115.6 mm in females), and B. spinata (54.0–55.0 mm in females). Boulenophrys puningensis sp. nov. shows the least genetic divergence from B. kuatunensis (mean p -distances 5.3 % in the 16S gene) and B. daiyunensis (mean p -distances 6.2 % in the 16S gene). However, the new species distinctively differs from these species by having relatively shorter shanks with the heels not meeting when the flexed hindlimbs are held at right angles to the body axis (vs. heels meeting or overlapping in B. daiyunensis); having rudiment of webbing and no lateral fringes on toes (vs. lateral fringes narrow in B. daiyunensis; webbing absent in B. kuatunensis); having raised and enlarged tubercles with spines on their tips on surface of posterior abdomen, ventral thighs and around the cloaca (vs. such tubercles not enlarged and without spines in B. daiyunensis; ventral surface smooth in B. kuatunensis). Boulenophrys puningensis sp. nov. is morphologically most similar to B. brachykolos, which is restricted to Hong Kong and Shenzhen, China (Liu et al. 2018). The new species differs from B. brachykolos by having vomerine teeth (vs. absent in B. brachykolos); lacking spines on the surface of the tympanic region (vs. having dense tiny spines on the surface of the tympanic region in B. brachykolos); and having different relative finger length formula (I = II Banophrys puningensis sp. nov. vs. II B. brachykolos). In having relatively shorter shanks with heels that do not meet when the flexed hind limbs are held at right angles to the body axis, Boulenophrys puningensis sp. nov. can be easily distinguished from the following 32 congeners, i.e. B. angka, B. anlongensis, B. baishanzuensis, B. baolongensis, B. binchuanensis, B. binlingensis, B. boettgeri, B. congjiangensis, B. cheni, B. chishuiensis, B. jiangi, B. jinggangensis, B. jiulianensis, B. leishanensis, B. lini, B. minor, B. mufumontana, B. nanlingensis, B. palpebralespinosa, B. qianbeiensis, B. sanmingensis, B. shimentaina, B. shunhuangensis, B. tongboensis, B. wuliangshanensis, B. wushanensis, B. xiangnanensis, B. xianjuensis, B. yaoshanensis, B. yangmingensis, B. yingdeensis and B. yunkaiensis, all of which have relatively longer shanks with the heels meeting or overlapping. By the presence of vomerine teeth, Boulenophrys puningensis sp. nov. differs from B. acuta, B. caobangensis, B. daoji, B. huangshanensis, B. lishuiensis, B. lushuiensis, B. obesa, B. ombrophila, B. tuberogranulatus, and B. wugongensis, all of which lack vomerine teeth. By having a rounded tongue margin that is not notched distally, Boulenophrys puningensis sp. nov. differs from B. hoanglienensis, and B. insularis, all of which have notched tongues. By the absence of lateral fringes on toes, Boulenophrys puningensis sp. nov. differs from B. rubrimera, which has narrow lateral fringes on toes. By the presence of rudimentary webbing on the toes, Boulenophrys puningensis sp. nov. differs from B. daweimontis, B. fansipanensis, and B. frigida, all of which lack webbing on the toes. Boulenophrys puningensis sp. nov. further differs from the remaining B. dongguanensis and B. nankunensis by having raised tubercles bearing spines on the surface of the posterior abdomen, ventral thighs, and around the cloaca (vs. absence of such tubercles and spines in B. dongguanensis and B. nankunensis). Description of holotype. Adult male. Body size small, SVL 34.6 mm. Head width slightly larger than head length, HWD/HDL 1.03; 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 0.36 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 large with an obvious margin, TD/ED 0.70; large ovoid choanae at base of maxilla; vomerine ridge and vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched distally; presence of single subgular vocal sac, and pair of slit-like openings at posterior of jaw. Forearm (i.e., radioulna) length 0.21 of SVL and hand 0.23 of SVL; hand without webbing, fingers without lateral fringes, relative finger length I = II Dorsal skin rough and granular, with raised conical tubercles; sparse large tubercles on flanks; single horn-like prominent tubercle on edge of upper eyelid; obvious supratympanic fold curving posteroventrally from posterior corner of eye to level above insertion of arm; tympanic region smooth without granules or tubercles; dense tubercles on skin of upper lip, upper eyelid, mandibular articulation, loreal, temporal region excluding tympanum and surface around cloaca; a single discontinuous “V” shaped ridge present on occipital region; dense tubercles on shanks and thighs; ventral surface with dense raised tubercles; tubercles on surface of posterior abdomen, ventral surface of thighs and around cloaca bearing tiny spines on their tips; small pectoral gland closer to axilla; single femoral gland positioned on posterior surface of thigh at midpoint between knee and cloaca. Coloration of holotype in life. Dorsal surface of body yellowish brown, with incomplete dark brown triangular marking between eyes. Two wide oblique black bands present on forearm. Dorsal surface of fingers and hindlimbs with dark grey transverse bands. Presence of vertical dark brown band below eye. Tubercles on edge of upper eyelid beige. Supratympanic fold light brown. Ventral surface dark grey, with black longitudinal band on surface of throat; surface of throat and chest mottled with orange patches. Tubercles on ventral surface of chest, belly, and thighs greyish white; spines on tips of tubercles on surface of posterior abdomen; ventral surface of thighs and around cloaca dark grey. Digits, inner and outer metacarpal tubercles and inner metatarsal tubercle greyish white. Pectoral glands and femoral glands beige, mottled with orange patches. Iris yellowish brown, with greyish white patches on upper and lower margin. Coloration of holotype in preservative. Yellowish brown fades to greyish brown dorsally. Color of the triangular marking between eyes, oblique bands on forearms, patterns on ventral surface faded. Orange patches on surface of throat, chest; color of pectoral glands and femoral glands faded. Variation. Mensural data of the type series are listed in Table 4. Most of the paratypes are similar to the holotype in morphology and color pattern, except for the following: dorsal surface of body yellowish brown in the holotype (vs. dorsal surface of body light brown in the paratypes SYS a007647 (Fig. 4C) and SYS a007648 (Fig. 4E); ventral surface dark grey with orange patches (vs. ventral surface lacking bright patches in the paratypes SYS a007649 (Fig. 4B), SYS a007647 (Fig. 4D) and SYS a007648 (Fig. 4F); iris yellowish brown, with greyish white patches on its upper and lower margin in the holotype (vs. iris grey with beige and dark mottling in the paratype SYS a007649 (Fig. 4A); tubercles on posterior part of abdomen of the paratype SYS a007648 (Fig. 4F) are weakly developed. Females are distinctly larger than the males. Distribution and natural history. Currently, Boulenophrys puningensis sp. nov. is only known from its type locality, Longkeng Village of Puning. It inhabits flowing montane streams and the nearby forest floor and leaf litter at elevations between 120– 300 m. Advertisement calls of males were heard from February until April. Males were found calling in rock crevices in the flowing streams. Tadpoles could be found in this period.

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.

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.

VIII. Incertae sedis with Megophryinae Two species: “ Megophrys ” dringi Inger, Stuebing & Tan, 1995; “ Megophrys ” feii Yang, Wang & Wang, 2018.

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.