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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.

Gekko (Japonicgekko) subpalmatus (Günther, 1864) Chresonymy. Gecko subpalmatus — Günther 1864. Gekko subpalmatus — Schmidt 1927 (part); Pope 1935 (part); Zhao et al. 1999 (part). Gekko (Japonigekko) subpalmatus — Wood et al. 2020. Type materials. Holotype. BMNH 1946.8.2592, adult female, collected by Fortune from Chikiang [= Zhejiang], PR China. Specimens examined. Eight adult male and three adult female specimens. Males SYS r001380–1381, collected from Donghai Canyon, Zhoushan City, Zhejiang Province, PR China; males SYS r001767 (Fig. 5A 1), 2065, 2068–2070, 2072, and females SYS r001762 (Fig. 5A 2), 1768, 2071, collected from Xikou Township, Fenghua District, Ningbo City, Zhejiang Province. Common name. Webbed-toed Gecko (in English); pǔ zhǐ bì hŭ (×flŧt). Etymology. The specific epithet subpalmatus is a compound adjective of sub- (‘under’, ‘beneath’ in Latin) and palmâtus (‘palmated’ in Latin), referring to the fingers and toes basally webbed in this species. Revised diagnosis. (1) medium-sized gecko species, SVL 50.9–65.8 mm in adults; (2) tubercles on dorsal body, limbs and tail absent; (3) eye moderate, ED/HL ratio 0.18–0.22; (4) rostral moderate, elongate transversely, RW/HW ratio 0.18–0.22, RW/RH ratio 2.08–2.30; (5) mental elongate transversely, MW/HW ratio 0.10–0.15, MW/ML ratio 1.00–1.70; (6) nares bordered by rostral, internasals 0–1; (7) interorbital scales between anterior corners of the eyes 28–37; (8) midbody scale rows 129–156; (9) ventral scale rows at midbody 39–46; (10) scales between mental and cloacal slit 144–190; (11) subdigital lamellae on first fingers 9–11, on fourth fingers 11–15, on first toes 9–12, on fourth toes 11–14; (12) fingers and toes with distinct webbing; (13) 5–9 precloacal pores in a continuous row in males; (14) a single postcloacal tubercle on both sides; (14) dorsum greyish white to dark brown, with five regular dark bands between nape and sacrum (Fig. 6A). Hemipenial characteristics (Fig. 3A): (1) hemipenis clavate, bilobed, densely covered with denticulate-seamed calyces; (2) sulcus spermaticus centrifugal, bifurcate at crotch of hemipenis; (3) lateral welt well developed, visible from asulcate side, not in contact with sulcus lip; (4) calyces confined to lobes and distal 2/3 of truncus; (5) tonguelike welt well-developed; (6) apical folds large, comma-shaped, in contact with each other; (7) a small and boomerang-like area on the asulcate side of the lobe center, calyces on this area not well developed (Fig. 3A, Ch. 7). Remark. The type locality of Gekko (Japonigekko) subpalmatus is in “Chikiang” [= Zhejiang] according to the original description, but the specific locus is unknown (Günther 1864; Pope 1935). Subsequently, this species has been reported from multiple localities in eastern, southern, and southeastern China (Schmidt 1927; Pope 1935; Zhao et al. 1999). Currently, G. (J.) subpalmatus can be recognized only from Zhejiang based upon voucher specimens and molecular data in this study. The populations in Fujian are temporally retained under the name of G. (J.) subpalmatus (see Discussion section). The previous records in eastern and southeastern Guangdong and southern Jiangxi should be as allocated to G. (J.) melli (Yang et al. 2012; this study), and the previous records in Sichuan, Guizhou, and Chongqing should be as allocated to G. (J.) cib sp. nov. (this study)., Published as part of Lyu, Zhi-Tong, Lin, Chao-Yu, Ren, Jin-Long, Jiang, Ke, Zhang, Yin-Peng, Qi, Shuo & Wang, Jian, 2021, Review of the Gekko (Japonigekko) subpalmatus complex (Squamata, Sauria Gekkonidae), with description of a new species from China, pp. 236-258 in Zootaxa 4951 (2) on page 247, DOI: 10.11646/zootaxa.4951.2.2, http://zenodo.org/record/4663976, {"references":["Gunther, A. C. L. G. (1864) The Reptiles of British India. Taylor & Francis, London, 452 pp., XXVII pls.","Schmidt, K. P. (1927) Notes on Chinese reptiles. Bulletin of the American Museum of Natural History, 54, 467 - 551","Pope, C. H. (1935) The Reptiles of China. Turtles, Crocodilians, Snakes, Lizards. Natural History of Central Asia. Vol. 10. American Museum of Natural History, New York, 604 pp.","Zhao, E., Zhao, K. & Zhou, K. (1999) Fauna Sinica. Reptilia vol. 2. Squamata (Lacertilia). Science Press, Beijing, 394 pp. [in Chinese]","Wood, P. L., Guo, X., Travers, S. L., Su, Y. C., Olson, K. V., Bauer, A. M., Grismer, L. L., Siler, C. D., Myle, R. G., Andersen, M. J. & Brown, R. M. (2020) Parachute geckos free fall into synonymy: Gekko phylogeny, and a new subgeneric classification, inferred from thousands of ultraconserved elements. Molecular Phylogenetics and Evolution, 146, 106731. https: // doi. org / 10.1016 / j. ympev. 2020.106731","Yang, J. H., Wang, Y. Y., Zhang, T. D., Sun, Y. J. & Lin, S. S. (2012) Genetic and morphological evidence on the species validity of Gekko melli Vogt, 1922 with notes on its diagnosis and range extension (Squamata: Gekkonidae). Zootaxa, 3505 (1), 67 - 74. https: // doi. org / 10.11646 / zootaxa. 3505.1.5"]}

Gekko (Japonicgekko) melli (Vogt, 1922) Chresonymy. Gecko melli — Vogt 1922 Gekko subpalmatus — Pope 1935 (part); Gressitt 1941; Bauer & Günther 1991; Zhao et al. 1999 (part). Gekko melli — Rösler et al. 2005; Rösler & Tiedemann 2007; Yang et al. 2012. Gekko (Japonigekko) melli — Wood et al. 2020 Type materials. Lectotype. ZMB 27659 A, adult male, collected by R. Mell from N. O. Kuangtung [= northeastern Guangdong], PR China; designated by Bauer & Günther (1991). Paralectotype. ZMB 27659 B, juvenile, same data as lectotype; designated by Bauer & Günther (1991). Specimens examined. Two adult male and five adult female specimens. Male SYS r000267 (Fig. 5B 1), collect- ed from Mt. Jiulian, Longnan County, Jiangxi Province, PR China; male SYS r000440, and females SYS r000438, 0439, 0451–0452, 0453 (Fig. 5B 2), collected from Mt. Yinping, Dongguan City, Guangdong Province, PR China. Common name. Mell’s Gecko (in English); meí shì bì hŭ (h氏ŧt). Etymology. The specific name melli is in reference to the collector Rudolf Emil Mell (1878–1970), who was the director of the German-Chinese Middle School at Canton [= Guangzhou City] and collected many animal specimens in Guangdong. His name appeared in Rösler & Tiedemann (2007) as “Robert Mell” due to a typographical mistake (Herbert Rösler, personal communication). Revised diagnosis. (1) medium-sized gecko species, SVL 64.5–80.3 in adults; (2) tubercles on dorsal body, limbs and tail absent; (3) eye moderate, ED/HL ratio 0.21–0.24; (4) rostral moderate, RW/HW ratio 0.20–0.22, RW/RH ratio 1.83–2.13; (5) mental elongate transversely, MW/HW ratio 0.14–0.18, MW/ML ratio 1.11–1.56; (6) nares bordered with rostral, internasals 1–2; (7) interorbital scales between anterior corners of the eyes 34–40; (8) midbody scale rows 148–160; (9) ventral scale rows at midbody 44–46; (10) scales between mental and cloacal slit 171–192; (11) subdigital lamellae on first fingers 9–11, on fourth fingers 9–14, on first toes 10–12, on fourth toes 11–14; (12) fingers and toes with distinct webbing; (13) 9–11 precloacal pores in a continuous row in males; (14) a single postcloacal tubercle on both sides; (14) dorsum greyish white to dark brown, with iregular large dark patches between nape and sacrum; (15) top of head with an small incomplete W-shaped marking, and posteriorly followed by a large inverted W-shaped marking on dorsal neck. (Fig. 6B). Hemipenial characteristics (Fig. 3B): (1) hemipenis clavate, bilobed, densely covered with denticulate-seamed calyces; (2) sulcus spermaticus centrifugal, bifurcate at half of truncus; (3) lateral welt developed, visible from asulcate side, in contact with sulcus lip; (4) calyces extend to lobes and proximal 1/3 of truncus; (5) tongue-like welt weakly developed; (6) apical folds small, arc-shaped, not in contact with each other; (7) a relatively large and rounded area on the asulcate side of the lobe, calyces on this area not well developed (Fig. 3B, Ch 7). Remark. The original description of Gekko (Japonigekko) melli in Vogt (1922) contained no information of locality or designation of type materials. The lectotype and paralectotype was designated by Bauer & Günther (1991) with providing the locality as “N. O. Kuangtung ” [= northeastern Guangdong] while the exact locus is unavailable. The lectotype and paralectotype were redescribed by Rösler & Tiedemann (2007). Currently, this species is recognized from multiple localities in eastern and northeastern Guangdong and neighboring southern Jiangxi (Yang et al. 2012; this study). G. (J.) melli is also expected to occur in southern Fujian which is geographically related to eastern Guangdong, but confirmation requires further vouchers (see Discussion section)., Published as part of Lyu, Zhi-Tong, Lin, Chao-Yu, Ren, Jin-Long, Jiang, Ke, Zhang, Yin-Peng, Qi, Shuo & Wang, Jian, 2021, Review of the Gekko (Japonigekko) subpalmatus complex (Squamata, Sauria Gekkonidae), with description of a new species from China, pp. 236-258 in Zootaxa 4951 (2) on pages 247-249, DOI: 10.11646/zootaxa.4951.2.2, http://zenodo.org/record/4663976, {"references":["Vogt, T. (1922) Zur Reptilien- und Amphibienfauna Sudchinas. Archiv fur Naturgeschichte, 88, 135 - 146.","Pope, C. H. (1935) The Reptiles of China. Turtles, Crocodilians, Snakes, Lizards. Natural History of Central Asia. Vol. 10. American Museum of Natural History, New York, 604 pp.","Gressitt, J. L. (1941) Amphibians and reptiles from southeastern China. The Philippine Journal of Science, 75, 1 - 58.","Bauer, A. M. & Gunther, R. (1991) An annotated type catalogue of the geckos (Reptilia: Gekkonidae) in the Zoological Museum, Berlin. Mitteilungen aus dem Zoologischen Museum in Berlin, 67, 279 - 310. https: // doi. org / 10.1002 / mmnz. 19910670204","Zhao, E., Zhao, K. & Zhou, K. (1999) Fauna Sinica. Reptilia vol. 2. Squamata (Lacertilia). Science Press, Beijing, 394 pp. [in Chinese]","Rosler, H., Ziegler, T., Vu, N. T., Herrmann, H. W. & Bohme, W. (2005 \" 2004 \") A new lizard of the genus Gekko Laurenti, 1768 (Squamata: Sauria: Gekkonidae) from the Phong Nha-Ke Bang National Park, Quang Binh Province, Vietnam. Bonner zoologische Beitrage, 53, 135 - 148.","Rosler, H. & Tiedemann, F. (2007) Gekko melli Vogt, 1922 and its types (Reptilia, Sauria, Gekkonidae). Mitteilungen aus dem Museum fur Naturkunde in Berlin, Zoologische Reihe, 83, 105 - 108. https: // doi. org / 10.1002 / mmnz. 200600033","Yang, J. H., Wang, Y. Y., Zhang, T. D., Sun, Y. J. & Lin, S. S. (2012) Genetic and morphological evidence on the species validity of Gekko melli Vogt, 1922 with notes on its diagnosis and range extension (Squamata: Gekkonidae). Zootaxa, 3505 (1), 67 - 74. https: // doi. org / 10.11646 / zootaxa. 3505.1.5","Wood, P. L., Guo, X., Travers, S. L., Su, Y. C., Olson, K. V., Bauer, A. M., Grismer, L. L., Siler, C. D., Myle, R. G., Andersen, M. J. & Brown, R. M. (2020) Parachute geckos free fall into synonymy: Gekko phylogeny, and a new subgeneric classification, inferred from thousands of ultraconserved elements. Molecular Phylogenetics and Evolution, 146, 106731. https: // doi. org / 10.1016 / j. ympev. 2020.106731"]}

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.

Gekko (Japonicgekko) cib sp. nov. Chresonymy. Gekko subpalmatus — Schmidt 1927 (part); Wu et al. 1985; Zhao et al. 1999 (part); Zhao 2003. Type materials. Holotype. CIB 116961 (Fig. 7), adult male, collected by Jin-Long Ren on 14 July 2018, from the wall surface in the Herpetological Museum, Chengdu Institute of Biology, Chinese Academy of Sciences (30.632093°N, 104.068917°E; ca 500 m a.s.l.), Chengdu City, Sichuan Province, PR China. Paratypes. Two adult male and eight adult female specimens. Females CIB 116962–116963 collected by Jin-Long Ren, Bo-Chen Zhao, and Zeng Wang on 28 June 2018 from Wenshu Temple (30.677950°N, 104.063181°E; ca 500 m a.s.l.), Chengdu City. Males SYS r002397–2398 and females SYS r002395–2396 collected by Zhi-Tong Lyu and Shuo Qi on 22 October 2019 from Chengdu City. Female SYS r000708 collected by Jian-Huan Yang on 5 June 2012 from Qingyinge, Mt. Emei (28.575478°N, 103.405444°E; ca 760 m a.s.l.), Emeishan City, Sichuan Province. Female SYS r001489 collected by Zhi-Tong Lyu on 16 June 2016 from Shunyangxi, Zihuai Township (28.628283°N, 106.298241°E; ca 800 m a.s.l.), Hejiang County, Sichuan Province. Females CIB 116964–116965 collected by Di-Hao Wu on 13 July 2017, from Shacuoluo, Malu Township (27.639252°N, 105.819409°E; ca 900 m a.s.l.), Jinsha County, Guizhou Province, PR China. Common name. CIB Gecko (“ CIB ” pronounced as “/si:/, /aɪ/, /bi:/”); chéng dű bì hŭ (成Ồ壁ẇ). Etymology. The specific name cib (pronounced as “/kib/”) is in reference to the abbreviation of the type locality, Chengdu Institute of Biology, Chinese Academy of Sciences (+afifks4±物ffṅNj). Chengdu Institute of Biology is a famous institute with a long and prestigious history in China, cultivating a large number of outstanding herpetologists and promoting the development of Chinese herpetology. Its English common name is directly derived from the scientific name. The Chinese common name is refers to the type locality of this species. Diagnosis. Gekko (Japonicgekko) cib sp. nov. is distinguished from all congeners by the following morphological characteristics: (1) medium-sized gecko species, SVL 46.9–66.4 mm in adults; (2) tubercles on dorsal body, limbs and tail absent; (3) eye large, ED/HL ratio 0.21–0.28; (4) rostral moderate, RW/HW ratio 0.19–0.24, RW/RH ratio 1.79–2.17; (5) mental elongate transversely, MW/HW ratio 0.13–0.18, MW/ML ratio 1.43–1.80; (6) nares bordered with rostral, internasals 1–2; (7) interorbital scales between anterior corners of the eyes 28–36; (8) midbody scale rows 128–149; (9) ventral scale rows at midbody 37–45; (10) scales between mental and cloacal slit 171–196; (11) subdigital lamellae on first fingers 8–13, on fourth fingers 10–15, on first toes 9–11, on fourth toes 10–15; (12) fingers and toes with distinct webbing; (13) 7–9 precloacal pores in a continuous row in males; (14) a single postcloacal tubercle on both sides; (14) dorsum greyish white to dark brown, with five regular dark bands between nape and sacrum (Fig. 6C). Hemipenial characteristics (Fig. 3C): (1) hemipenis clavate, bilobed, densely covered with denticulate-seamed calyces; (2) sulcus spermaticus bifurcate at half of truncus; (3) lateral welt weakly developed, invisible from asulcate side, in contact with sulcus lip; (4) calyces confined to lobes and distal 1/2 of truncus; (5) tongue-like welt large; (6) apical folds large, comma-shaped, not in contact with each other; (7) a small and boomerang-like area on the asulcate side of the lobe center, calyces on this area not well developed (Fig. 3C, Ch 7). Description of holotype. CIB 116961 (Fig. 7), an adult male with a total length of 98.8 mm (SVL 46.9 mm, TaL 51.9 mm); body slender, elongate (AG/SVL 0.47); head longer than wide (HL/HW 1.24); rostral quadrangular without suture medially, nearly twice wider than high (RW/RH 2.10) and wider than mental (RW/MW 1.50), touching first supralabial and supranasal on each side; nostrils oval, in contact with rostral, first supralabial, supranasal, and two enlarged nasals posteriorly, upper nasal smaller than lower nasal; posterior nasal region concave; internasals 2, slightly enlarged; preorbitals 15/16, preorbital region deeply concave; interorbitals 31; eye large (ED/HL 0.25), pupil vertical, elliptic; ear opening oval, horizontal, smaller than eye (TD/ED 0.34); mental trapezoid, about two times wider than long (MW/ML 1.75); postmental single, hexagonal, length about equal to width, shorter than length of mental, touching mental and first infralabial on both sides and 4 gular scales posteriorly, gular scales about equal size; supralabials 11/13; infralabials 13/13; dorsal scales on body smooth, round or oval, granular and juxtaposed, tubercles absent; lateral fold present; ventrals distinctly larger than dorsal scales, smooth, imbricate, and largest in the middle of belly; ventral scale rows at midbody 40; scale rows around midbody 135; ventral scales in a row between mental and cloacal slit 180; scales on dorsal forelimbs slightly enlarged; no tubercles on dorsal surface of limbs; scales on anterior and ventral parts of femur larger than those on posterior and dorsal parts; fingers and toes basally webbed; subdigital lamellae under first finger 9/9, under fourth finger 11/10, under first toe 10/10, under fourth toe 11/12; precloacal pores 8, in a continuous row; 11 precloacal scales rows and distinctly smaller posteriorly, three rows after precloacal pores enlarged; postcloacal tubercles 1/1; base of tail thickened, without tubercles on dorsal surface; subcaudals in a single median series, smooth, imbricate. Coloration of holotype. In life, dorsal surface of head brownish grey, speckled with dark, grey, and beige; labials yellowish, with grey bars; upper eyelids brownish yellow; dorsal surface of body brownish grey with black and pale grey blotches, several enlarged pale grey blotches present along the vertebral column, some of them bordered with black anteriorly; dorsal surface of neck with an indistinct W-shaped marking; dorsal surface of fore and hind limbs pale grey, finely speckled with darker spots, outer region tending to beige especially on fingers and toes; ventral surface of head, belly, and limbs cream with small black spots; dorsal surface of tail with eleven grey transverse bands, becoming darker and more distinct posteriorly; ventral tail pale grey in forepart and with nearly closed bands distally. In preservative (Fig. 7), the pattern of recently preserved specimens resembles that of live animals, but the pale grey coloration on the dorsal surface of body and limbs becomes darker. Variations. Measurements and scale counts of type series specimens are given in Table 5. Ground color on dorsal surface of head, body and tail is also different among each individual from yellowish grey to blackish grey in the wild, most individuals body color become darker after capture. Hemipenial morphology. Description based on paratype SYS r002398 (Fig. 3C). Right hemipenis, fully evert- ed organ 6.00 mm long, 4.48 mm wide, stout, clavate, shallowly bilobed; densely covered with small or tiny calyces, seams of calyces denticulated. Two lobes subequal, both the lobes covered with calyces, calyces on distal part of lobes tinier; calyces proximally extend to the upper half part of truncus, bordered by slightly protruded fold. Each lobe with a distinctly protruded, comma-like apical folds, two folds not in contact with each other. On sulcal side, a distinct tongue-like welt present at crotch, curled inward. A thin, wrinkled, lateral welt present on the left side of sulcal lip, in contact with sulcal lip at distal one third of truncus. On asulcate side, lateral welt invisible, a small and boomerang-like area present on the center of each lobe, calyces on this area not well developed. Sulcus spermaticus forked, bordered by voluminous skin bulges, bifurcate at middle part of truncus, terminating on the upper side of each lobe in a slit-like concavity. sulcal lips moderately protruded, gradually become weaker and thinner towards the distal part of the organ. Comparisons. Gekko (Japonigekko) cib sp. nov. is compared with all 30 recognized species within the subgenus Japonigekko. Gekko (Japonigekko) cib sp. nov. used to be recorded as G. (J.) subpalmatus. However, the new species can be distinguished from the true G. (J.) subpalmatus in eastern China by the following morphological characters: mentals more elongate transversely, MW/ML ratio 1.60 ± 0.12 (versus 1.36 ± 0.18); larger eyes, ED/HL ratio 0.23 ± 0.03 (versus 0.19 ± 0.01). In addition, G. (J.) cib sp. nov. differs from G. (J.) subpalmatus in the following distinct hemipenial characters (Table 3; Fig. 3), including having sulcus spermaticus bifurcate at half of the truncus length (vs. bifurcate at crotch of hemipenis), lateral welt of the base of the sulcus spermaticus weakly developed, in contact with sulcus lip (versus well developed, not in contact with sulcus lip), and a smaller ornamented area (calyces confined to lobes and distal 1/2 of truncus versus calyces extended to lobes and distal 2/3 of truncus). Gekko (Japonigekko) cib sp. nov. is the sister taxon to G. (J.) melli from southern China, however, it differs in the appearance, dorsal body with regular patches (versus dorsal body with irregular enlarge patches), absence of an incomplete W-shaped marking on top of head (versus present), W-shaped marking on dorsal neck narrow (versus W-shaped marking on dorsal neck broad), and in the following data of scale counts: (1) fewer scales around the middle of the body, SR 128–149 (versus SR 147–160), (2) fewer precloacal pores in males, PP 7–9 (versus PP 9–11). Particularly, G. (J.) cib sp. nov. differs from G. (J.) melli in the following distinct hemipenial characters (Table 3; Fig. 3), including having lateral welt of the base of the sulcus spermaticus weakly developed, invisible from asulcate side (versus developed, visible from asulcate side); developed tongue-like welt (versus weakly developed); a smaller ornamented area (calyces confined to lobes and distal 1/2 of truncus versus calyces extended to lobes and distal 2/3 of truncus); and a protruded boundary between naked area and ornamented area on truncus (versus boundary smooth). ......continued on the next page Gekko (Japonigekko) cib sp. nov. can be easily distinguished from the following 19 congeners by lacking dorsal tubercles: G. (J.) adleri Nguyen, Wang, Yang, Lehmann, Le, Ziegler & Bonkowski, 2013, G. (J.) auriverrucosus Zhou & Liu, 1982, G. (J.) canhi Rösler, Nguyen, Van Doan, Ho, Nguyen & Ziegler, 2010, G. (J.) chinensis (Gray, 1842), G. (J.) hekouensis Pope, 1928, G. (J.) japonicus (Schlegel, 1836), G. (J.) kwangsiensis Yang, 2015, G. (J.) lauhachindai Panitvong, Sumontha, Konlek & Kunya, 2010, G. (J.) liboensis Zhou, Liu & Li, 1982, G. (J.) pal- matus Boulenger, 1907, G. (J.) scabridus Liu & Zhou, 1982, G. (J.) shibatai Toda, Sengoku, Hikida & Ota, 2008, G. (J.) similignum Smith, 1923, G. (J.) swinhonis Günther, 1864, G. (J.) taibaiensis Song, 1985, G. (J.) vertebralis Toda, Sengoku, Hikida & Ota, 2008, G. (J.) vietnamensis Sang, 2010, G. (J.) wenxianensis Zhou & Wang, 2008, and G. (J.) yakuensis Matsui & Okada, 1968. Furthermore, the new species can be distinguished from the following three Gekko (Japonigekko) congeners by having only a single postcloacal tubercle on both sides (versus PAT 2– 2 in G. (J.) sengchanthavongi Luu, Calame, Nguyen, Le & Ziegler, 2015, and G. (J.) thakhekensis Luu, Calame, Nguyen, Le, Bonkowski & Ziegler, 2014; PAT 2–3 in G. (J.) scientiadventura Rösler, Ziegler, Vu, Herrmann & Böhme, 2004). For the remaining six subgeneric congeners, by having 7–9 precloacal pores in males, Gekko (Japonigekko) cib sp. nov. can be distinguished from G. (J.) aaronbaueri Tri, Thai, Phimvohan, David & Teynié, 2015 (PP 3–4), G. (J.) bonkowskii Luu, Calame, Nguyen, Le & Ziegler, 2015 (PP 6), G. (J.) nadenensis Luu, Nguyen, Le, Bonkowski & Ziegler, 2017 (PP 6), G. (J.) tawaensis Okada, 1956 (PP 0), and G. (J.) truongi Phung & Ziegler, 2011 (PP 10–11); by having 1–2 intersupranasals, the new species can be distinguished from G. (J.) bonkowskii and G. (J.) nadenensis, both of which have no intersupranasals; and by having 5 dark bands between nape and sacrum on dorsum, G. (J.) cib sp. nov. further differs from G. (J.) guishanicus Lin & Yao, 2016 (dorsum with 6 dark bands). Distribution and ecology. Currently, Gekko (Japonigekko) cib sp. nov. is recognized from Chengdu City, Mt. Emei, and Hejiang County of Sichuan, and Jinsha County from Guizhou. These localities all situated in or around the Sichuan Basin, suggesting the previous records of G. (J.) subpalmatus from these regions (including Sichuan, Guizhou, and Chongqing) should be reassigned to this new species. All individuals were found on house walls or under eaves.

Oligodon bivirgatus sp. nov. Figures 2–5 urn:lsid:zoobank.org:act: CDBD6461-6F4E-46F5-A65D-8E10AD89057D Holotype: SYNU 1907027, adult female, collected on 13 July 2019 by Tianyu Qian and Minghong Huang from Shangxi Nature Reserve (18.834167° N, 110.130833° E; 319 m a.s.l.), Hainan Province, China. Paratype: SYNU 1808001, adult female, collected on 17 August 2018 by Pipeng Li, Tianyu Qian and Minghong Huang from the same valley as the holotype (18.833611° N, 110.130556° E; 317 m a.s.l.). Etymology. The specific name comes from Latin words “ bi -” (double) and “ virgatus ” (striped), refers to this species has a dorsal color pattern made of two distinct dorsal stripes. The common name is recommended to read “Double Striped Kukri Snake” in English while it is “ Aäṁn'ḃ (Shuāng Xiàn Xiǎo Tóu Shé)” in Chinese. Diagnosis. Oligodon bivirgatus sp. nov. can be distinguished from its congeners known to occur in Southern China and mainland Southeast Asia by having the following combination of characters: (1) eight maxillary teeth; (2) nasal divided; (3) absence of a loreal; (4) single preocular and two postoculars; (5) seven supralabials, third and fourth touching the orbit, sixth excluded from the lip; (6) seven infralabials, anterior four contacting chin shields; (7) internasals separate from prefrontals; (8) single anterior and two posterior temporals; (9) anal plate divided; (10) 15/15/15 dorsal scale rows; (11) dorsum reddish-brown in life, with two distinct narrow dark stripes, two dorsolateral lines are faintly visible; and (12) ventral surface reddish in life, edge cream with irregular dark blotches. Description of holotype. Body slender, TL 465 mm, TaL 61 mm, SVL 404 mm; tail robust, tapering and quite short, Tal/TL ratio 13.1%; head short, HL 16.8 mm, HW 9 mm, slightly flattened, HH 6.3 mm, neck moderately marked; snout elongate, SnL 5.1 mm, SnL/HL ratio 30%. Head scalation. Details of head scalation are shown in Figure 3. Head scalation includes single rostral, two internasals, two prefrontals, two supraoculars, single frontal, and two parietals from dorsal view. Rostral shield large, thick, in contact with first supralabial, nasal and internasals posteriorly, curved onto upper snout surface, well visible from above; internasals in narrow contact, separated by a median suture; prefrontals sub-rectangular, width greater than length, separated by a suture; suture between internasals about equal to that between prefrontals; frontal large, pentagonal, posterior portion pointed, contacting parietal; supraoculars sub-rectangular, length greater than width distinctly, anterior narrower; parietals large, suture between parietals slightly shorter than length of frontal; nasal vertically divided, posterior part distinctly smaller, nostrils open in anterior part of nasal; loreal absent; pupil round, ED 2.2 mm, IO 5.4 mm; 1/1 vertically elongate, rectangular preocular; 2/2 rectangular postoculars; 1/1 anterior temporals, sub-rectangular, twice longer than wide, 2/2 posterior temporals, upper one smaller; 2 pair of chin shields, anterior pair elongate, the length is one third of the width, posterior pair sub-rectangular; SL 7/7, 1 st and 2 nd contacting nasal, 2 nd contacting internasal, 2 nd and 3 rd contacting preocular, 3 rd and 4 th contacting orbit, 4 th and 5 th contacting lower postocular, 6 th excluded from the lip, 5 th and 7 th barely distinguishable in size; IL 7/7, 1 st pair in contact with each other, 1 st– 4 th contact with anterior chin shields, 4 th largest, touching posterior chin shields. Body scalation. Dorsal scale rows 15/15/15, all smooth; vertebral scales similar to other dorsal scales in shape and size; ventral scales 173, angulate laterally; anal plate divided; subcaudals 37, all paired, terminal caudal scale forming a tip. Coloration of holotype. In life: Dorsal surface of head reddish-brown as dorsum, but paler on snout and rostral. Three pointing forward chevrons on head and neck. Chevrons maroon, somewhat paler maroon in the mesial, edged by darker pigmentation. The first chevron with its apex at the level of the posterior edge of the rostral, covering part of internasals, prefrontals, supraoculars and frontal, and descending through the eye to cover 3 rd– 5 th SL. The apex of the second chevron starts at the middle of head and covers the posterior part of frontal, extending to neck on both sides of the head, through parietals, upper anterior temporals, two posterior temporals, and descending behind the head, covering part of 7 th SL and 7 th IL. The third chevron, with its anterior tip on the suture of parietals is connected to the second chevron in the middle, forming an arrow, this arrow gradually widens and extends from parietals to the nape. The two edges of this arrow extend to dorsal as two narrow dark stripes, ending at the tip of tail. At the beginning of dorsal stripes, there are small blotches at the lateral of neck, about 3 rows scale rows behind head. The dorsal stripes are about 1/3 scale wide, covering 6 th and 10 th scale rows; another two dashed lines are faintly visible on the flanks. Ventral surface of head cream with black spots on the anterior portion of 1 st, 3 rd– 6 th SL, anterior chin-shields, other small spots at the midline of the gular scales. Belly is coral-red with its edge cream and black blotches irregularly distributed on each side. The blotches are small on the body, at intervals of one or two ventral scales, sometimes continuous, 95 on left and 100 on right. Ventral surface of the tail shows the same pattern as the belly. On the tail, the blotches become denser and no more intervals at the end 10 pairs of subcaudals, 27 on left and 25 on right. Iris bicolored, upper part light yellow, lower part brown, matching the dorsal pattern; pupil round, black with golden edge. In preservative: The dorsal color became paler and, eventually, totally grey. The head mark and the dorsum stripe remained maroon. The coral belly faded to white, black blotches on belly lightened and, as a consequence, reduced in size. Skull. The description of the skull of Oligodon bivirgatus sp. nov. is based on the three dimensional (3D) reconstructed model of the holotype (SYNU 1907027, Figure 5) and paratype (SYNU 1808001). Dorsally, the skull of Oligodon bivirgatus sp. nov. is roughly oval in shape. Nasal bones are slender and triangularly shaped, bilateral nasal bones fused at the posterior 2/3 part. The anterior part of the maxilla (in front of the ectopterygoid process) presents as lamella shaped. Dorsal-lateral ridge conspicuously ranges throughout the maxilla. Maxilla bears eight maxillary teeth on each side: two small normal teeth and six blade-like teeth, increasing in size posteriorly (the rearmost one largest), not separated by any diastema. The cutting edge of the blade-like teeth bulges posteriorly. Thus, the blade teeth look quite similar to the kukri knives. In contrast to most colubrid snakes, a small empty tooth position (one diastema in size) presents on the rostral tip of the ventral edge of the maxilla, which forms a small round-cornered process with the anterior part of the maxilla. Septomaxilla quite wide and flat. The conchal process of septomaxilla slightly curved posteriorly. Vomer quite robust and short. The anterior tip of vomer is rounded and bunt. The ascending process of premaxilla is quite short and considerably slanted anteriorly, barely contacting the nasals near the rostral bottom tip of the vertical laminae. The palatine is lamellar, the choanal process of palatine is quite slender, round cornered and ventrally curled. The maxilla process of palatine is round and bunt, slightly ventrally curved. One section of small vacuity (approximately two palatine teeth in distance) presents at the tip of the ventral edge of the rostral, which form a small round-corner process in the anterior part of palatine. Ectopterygoid quite short, thick and robust, the angular surface to the maxilla is spanner-shaped. Postorbital slender, rostrally curved and elongated, not contacting the frontal. Supratemporal bone is slender and triangular; the quadrate-connecting surface takes up about 3/4 the length of the total supratemporal. The anterior and posterior tip are slightly sharp in shape. Quadrate bone triangular, roughly perpendicular to the mandible, the angular surface to the supratemporal is quite wide and the one to mandible is thick and robust. Mandible: Compound bone quite slender and prominently curved ventrally. A prearticular crest is prominent and rounded. Two large oval foramina nutricia on the labial side of the dental bone. Dentition. Maxillary teeth 8/8 (2+6); palatine teeth 4/4 (3/4); dentary teeth 13/11 (11/10); pterygoid teeth: asymmetry in number, 9/4 (8/4), degenerated in size, significantly smaller than the ones on the palatomaxillary arch and dental bones. Variation. Morphological traits and pholidosis of two type specimens are listed in Table 4. The paratype matches most characters of the holotype, but there are little differences between them. Scalation. The internasals are not in contact with the posterior part of nasal in holotype (SYNU 1907027), but contacting in the paratype (SYNU 1808001). ......continued on the next page Coloration. There are two differences in dorsal coloration on head between the paratype (SYNU 1808001) and the holotype (SYNU 1907027): 1. (Paratype) The tip of the second chevron on head is in the middle of frontal and in contact with first chevron, a gap is formed in back of the middle, but no contact and divided posterior portion in holotype; 2. (Paratype) The third chevron commences at the posterior end of parietals with a blunt tip. The condition of the third chevron in the holotype differs in that it commences at the middle of the parietals and forms a sharp angle. Comparisons. Oligodon bivirgatus sp. nov. share similar scale patterns with O. ornatus, but differs from the latter by having different dorsal coloration (longitudinal stripes vs. crossbars in O. ornatus), and ventral coloration (wide red band vs. narrow red line in O. ornatus). The dorsal coloration of O. bivirgatus is also similar with O. pseudotaeniatus, but this new species can be distinguished from the latter by having 15/15/15 dorsal scale rows (vs. 17/17/ 15 in O. pseudotaeniatus), a divided anal plate (vs. entire in O. pseudotaeniatus), absence of a loreal (vs. present in O. pseudotaeniatus), a larger count of 173–176 ventrals (vs. 137–156 in O. pseudotaeniatus), and smaller count of 8 maxillary teeth (vs. 15 in O. pseudotaeniatus). Additional morphological comparative data of Oligodon bivirgatus sp. nov. and 41 recognized Oligodon species occurred in mainland Southeast Asia are shown in Table 5. By having 15/15/15 dorsal scale rows, Oligodon bivirgatus sp. nov. can be distinguished from its congeners occurring in mainland Southeast Asia having different numbers of MSR: O. albocinctus (19), O. annamensis (13), O. barroni (17), O. catenatus (13), O. cattienensis (17), O. chinensis (17), O. cinereus (17), O. condaoensis (17), O. cruentatus (17), O. culaochamensis (17 or 19), O. deuvei (17), O. eberhardti (13), O. fasciolatus (21 or 23), O. formosanus (19 or 17), O. joynsoni (17), O. lipipengi (19), O. macrurus (17), O. mcdougalli (13), O. melanozonatus (17), O. moricei (17), O. mouhoti (17), O. nagao (17 or 15), O. ocellatus (19), O. pallidocinctus (17), O. planiceps (13), O. saintgironsi (17 or 18), and O. taeniatus (19). By having a divided nasal, Oligodon bivirgatus sp. nov. can be distinguished from its congeners occurring in mainland Southeast Asia having an entire nasal scale: O. catenatus, O. cattienensis, O. eberhardti, O. erythrorhachis, O. hamptoni, O. lacroixi, O. lungshenensis, O. melanozonatus, O. planiceps, and O. torquatus. By having a divided anal plate, Oligodon bivirgatus sp. nov. can be distinguished from its congeners occurring in mainland Southeast Asia having an entire anal plate: O. albocinctus, O. annamensis, O. barroni, O. cattienensis, O. chinensis, O. cinereus, O. condaoensis, O. culaochamensis, O. deuvei, O. fasciolatus, O. formosanus, O. inornatus, O. joynsoni, O. kampucheaensis, O. lipipengi, O. macrurus, O. moricei, O. mouhoti, O. nagao, O. ocellatus, and O. pallidocinctus. By the absence of a loreal, Oligodon bivirgatus sp. nov. can be distinguished from its congeners occurring in mainland Southeast Asia having one loreal or two: O. albocinctus, O. barroni, O. cattienensis, O. chinensis, O. cinereus, O. condaoensis, O. cruentatus, O. culaochamensis, O. deuvei, O. dorsalis, O. eberhardti, O. fasciolatus, O. formosanus, O. inornatus, O. jintakunei, O. joynsoni, O. kampucheaensis, O. lipipengi, O. melaneus, O. moricei, O. mouhoti, O. nagao, O. ocellatus, O. pallidocinctus, O. planiceps, O. saintgironsi, and O. torquatus. By having eight maxillary teeth, Oligodon bivirgatus sp. nov. can be distinguished from its congeners occurred in mainland Southeast Asia having a larger or smaller number of maxillary teeth: O. albocinctus (10–12), O. barroni (10–13), O. chinensis (9–10), O. cinereus (10–12), O. condaoensis (11–13), O. cruentatus (14–16), O. culaochamensis (9–10), O. deuvei (12–15), O. fasciolatus (9–10), O. formosanus (10–11), O. inornatus (10–11), O. joynsoni (11–12), O. kampucheaensis (11), O. lipipengi (11), O. macrurus (13), O. melaneus (7), O. moricei (12), O. mouhoti (14–16), O. nagao (9–10), O. ocellatus (10), O. pallidocinctus (10–12), O. planiceps (10), O. pseudotaeniatus (15), O. rostralis (6), O. saintgironsi (10–12), O. taeniatus (14–17), and O. torquatus (15–16). By having a dorsal coloration of longitudinal stripes, Oligodon bivirgatus sp. nov. differs from its congeners occurred in mainland Southeast Asia having a dorsal coloration of crossbars or blotches: O. albocinctus, O. annamensis, O. barroni, O. cattienensis, O. chinensis, O. culaochamensis, O. erythrorhachis, O. fasciolatus, O. formosanus, O. jintakunei, O. joynsoni, O. kampucheaensis, O. lipipengi, O. melanozonatus, O. nagao, O. ocellatus, O. pallidocinctus, O. rostralis, and O. saintgironsi. Distribution and natural history. The new species is currently known only from Shangxi NR, Hainan, China. Both specimens were found at the same site in a stream of tropical lowland evergreen forest (Figure 6A). Snakes were observed about 19:30 hr–20:00 hr at night, so we believe that Oligodon bivirgatus sp. nov. is nocturnal. The paratype (SYNU 1808001) was found swimming in the slow stream, but crawled quickly when disturbed. The holotype (SYNU 1907027) was found on rock bank covered with moss along the stream (Figure 6B). O. bivirgatus sp. nov. are found to be sympatric with its congener O. formosanus, while the latter are larger in body size and more common in Shangxi NR. The other sympatric reptile species include Hebius boulengeri (Gressitt, 1937), Bungarus multicinctus Blyth, 1861, Trimerodytes percarinatus (Boulenger, 1899), Goniurosaurus hainanensis Barbour, 1908, Tropidophorus hainanus Smith, 1923, Sphenomorphus incognitus (Thompson, 1912) and Sphenomorphus tonkinensis Nguyen, Schmitz, Nguyen, Orlov, Böhme and Ziegler, 2011.

Zhou, Zhengyan, Sun, Zhiyong, Qi, Shuo, Lu, Yuyan, Lyu, Zhitong, Wang, Yingyong, Li, Pipeng, Ma, Jianzhang (2019): A new species of the genus Hebius (Squamata: Colubridae: Natricinae) from Hunan Province, China. Zootaxa 4674 (1): 68-82, DOI: https://doi.org/10.11646/zootaxa.4674.1.3