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The genus Leptobrachella is a species-rich genus of megophrid frog. Rapid discovery of many new species within this genus emphasizes the importance of regional research. In this study, we describe a new species of Leptobrachella, Leptobrachella verrucosasp. nov., from northwestern Guangdong Province, China, based on a combination of molecular and morphological data. A key including congeners from the same province, namely L. laui, L. liui, L. mangshanensis, L. shimentaina, and L. yunkaiensis, is provided.

A new species of Leaf Litter Toad,Leptobrachella shimentainasp. nov., is described from the Shimentai and Luokeng nature reserves of northern Guangdong Province, southern China. The new taxon can be distinguished from all recognized congeners by a combination of discrete morphological character state differences relating to its small body size (SVL 26.4–28.9 mm in six adult males, 30.1 and 30.7 mm in two adult females); a number of apparently fixed color pattern character differences (including eye coloration and color pattern features from dorsal, ventral, and dorsolateral surfaces of its head, body, limbs, and ventrum); the morphological and discrete characteristics of the external phenotype (the skin texture of dorsum and ventrum, the presence of supra-axillary and ventrolateral glands, the wide dermal fringes and rudimentary webbing on toes, and the uninterrupted longitudinal ridges under toes). Two samples of this new species previously were proposed as representing a new, unnamed species. We now substantiate this claim by providing diagnostic comparisons of discrete character differences. In addition, we also discuss taxonomic uncertainty surrounding the identity of two congeners,L. chishuiensisandL. purpurus, which we interpret as indicative of taxonomic inflation in the species-rich subfamily Megophryidae.

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.

The diversity of Asian horned toads is considered highly underestimated and to contain a large number of undescribed cryptic species. In this work, we describe three new species of Boulenophrys from south China, namely, Boulenophrys yaoshanensis sp. nov. from central Guangxi, Boulenophrys yingdeensis sp. nov. from northern Guangdong, and Boulenophrys yunkaiensis sp. nov. from western Guangdong. These three new species can be distinguished from all recognized congeners by a combination of morphological characteristics and significant genetic divergences. These descriptions increase the number of recognized species of Boulenophrys to 61. In addition, an updated checklist of the Asian horned toads of the subfamily Megophryinae is provided in this study.

Karstic landscapes play an important role in biodiversity formation and often contain high levels of endemism. However, site-endemic taxa in karstic landscapes are being threatened by exploitation and weak legal protection. In this study, we describe Odorrana concelata Wang, Zeng, & Lin, sp. nov., a limestone karst-restricted odorous frog from northern Guangdong, China. This new species shows distinctive genetic divergence and morphological differences from its congeners. Phylogenetic results suggest that the new species represents an independent lineage that is grouped with O. lipuensis and O. liboensis based on the mitochondrial 16S and 12S ribosomal RNA genes. We recommend the new species be listed as Vulnerable (VU) in the IUCN categorization as it is only known from the type locality with limited microhabitats and is threatened by habitat degradation.

A new species of colubrid snake,Lycodon cathayasp. nov., is described based on two adult male specimens collected from Huaping Nature Reserve, Guangxi, southern China. In a phylogenetic analyses, the new species is shown to be a sister taxon to the clade composed ofL. futsingensisandL. namdongensiswith low statistical support, and can be distinguished from all known congeners by the significant genetic divergence in the mitochondrial cytochromebgene fragment (p-distance ≥ 7.9%), and morphologically by the following combination of characters: (1) dorsal scales in 17–17–15 rows, smooth throughout; (2) supralabials eight, third to fifth in contact with eye, infralabials nine; (3) ventral scales 199–200 (plus two preventral scales), subcaudals 78; (4) loreal single, elongated, in contact with eye or not, not in contact with internasals; (5) a single preocular not in contact with frontal, supraocular in contact with prefrontal, two postoculars; (6) maxillary teeth 10 (4+2+2+2); (7) two anterior temporals, three posterior temporals; (8) precloacal plate entire; (9) ground color from head to tail brownish black, with 31–35 dusty rose bands on body trunk, 13–16 on tail; (10) bands in 1–2 vertebral scales broad in minimum width; (11) bands separate ground color into brownish black ellipse patches arranged in a row along the top of body and tail; (12) elliptical patches in 3–6 scales of the vertebral row in maximum width; (13) ventral surface of body with wide brownish black strip, margined with a pair of continuous narrow greyish white ventrolateral lines. With the description of the new species, 64 congeners are currently known in the genusLycodon, with 16 species occurring in China.

Recent phylogenetic analysis encompassing multilocus nuclear-gene and matrilineal mtDNA genealogy has revealed a series of cryptic species of the subgenus Panophrys within genus Megophrys from southern and eastern China. This study demonstrates that the Panophrys specimens from the hilly areas among Guangdong, Guangxi and Hunan can be morphologically distinguished from all recognized congeners, thereby providing additional supports for the recognitions of four new species of Panophrys, namely Megophrys (Panophrys) mirabilis Lyu, Wang & Zhao, sp. nov. from northeastern Guangxi, Megophrys (Panophrys) shimentaina Lyu, Liu & Wang, sp. nov. from northern Guangdong, and Megophrys (Panophrys) xiangnanensis Lyu, Zeng & Wang, sp. nov. and Megophrys (Panophrys) yangmingensis Lyu, Zeng & Wang, sp. nov. from southern Hunan. The descriptions of these species take the number of Megophrys species to 101, 46 of which belong to the subgenus Panophrys.

The taxonomic status of the previous record of Opisthotropis maculosa Stuart & Chuaynkern, 2007 from Guangdong and Guangxi, southern China, is revised based on the comparison of morphological and molecular data collected from the Chinese specimens and the holotype of O. maculosa from Thailand and O. haihaensis Ziegler, Pham, Nguyen, Nguyen, Wang, Wang, Stuart & Le, 2019 from Vietnam. Results reveal that the population from Shiwandashan Nature Reserve in southern Guangxi, China belongs to O. haihaensis, and represents the first national record for China; the populations from western Guangdong and southeastern Guangxi are described as a new species, Opisthotropis hungtaisp. nov. We suggest that O. maculosa should be removed from the Chinese herpetofauna checklist. The new national record of O. haihaensis and the description of the new species bring the total number of Opisthotropis to 13 in China.

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 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"]}

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 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."]}

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

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

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

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

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

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

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

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

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

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

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.

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

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

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

The Amolops populations in the coastal hills in eastern Guangdong and southern Fujian, China, were controversially recorded as A. hongkongensis or A. daiyunensis before. In this study, based on the morphological examination and phylogenetic analysis of the specimens from these areas, a new species, Amolops teochew sp. nov., is described. Amolops teochew sp. nov. can be distinguished reliably from A. hongkongensis and A. daiyunensis by a combination of characteristics morphologically and distinct divergences genetically. The description of the new species highlights the Amolops diversity in the limited hilly region of southeastern China, which is remarkably higher than that in the more extensive inland region of southeastern China.

Panophrys sanmingensis Lyu & Wang sp. nov. Chresonymy. Megophrys sp15— Liu et al. 2018 Holotype. SYS a002498 (Fig. 5 A–D), adult male, collected by Ying-Yong Wang on 18 April 2014 from Mt Longqi (26.5233°N, 117.2976°E; ca 700 m a.s.l.), Jiangle County, Sanming City, Fujian Province, PR China. Paratypes. Five adult males (SYS a002493–2496, SYS a002499/ CIB 116077) and one adult female (SYS a002500), collected at the same time from the same locality as the holotype. Etymology. The specific epithet sanmingensis refers to the type locality of the new species, the Sanming City. Common names. Sanming Horned Toad (in English) / Sān Míng Jiǎo Chán (Ξfifflḃin Chinese) Diagnosis. (1) body size small, with SVL 27.0– 29.5 mm (n = 6) in adult males and 29.5 mm in adult female; (2) tympanum distinct, TD/ED 0.49–0.50; (3) vomerine teeth absent; (4) margin of tongue notched; (5) heels overlapping; (6) tibio-tarsal articulation reaching just posterior to eye; (7) TIB/SVL 0.44–0.48, FTL/SVL 0.61–0.65; (8) narrow lateral fringes on fingers present, one subarticular tubercle present at the base of each finger, relative finger lengths I I in breeding males. Comparisons. Comparative data of Panophrys sanmingensis sp. nov. with P. daiyunensis, P. daoji, and 47 recognized Panophrys congeners are given in Table 3. Panophrys sanmingensis sp. nov. differs from P. daiyunensis by its vomerine teeth absent (vs present), tongue notched (vs not notched), lateral fringes on toes wide (vs narrow), and nuptial spines present on the finger I in breeding males (vs absent). Panophrys sanmingensis sp. nov. differs from P. daoji by its smaller body size, SVL 27.0– 29.5 mm in males and 29.5 mm in female (vs SVL 32.6–33.6 mm in males and 37.5–41.4 mm in females), tongue notched (vs not notched), heels overlapping (vs not meeting), and lateral fringes on toes wide (vs narrow). Compared with the six Panophrys species previously recorded from eastern China, P. sanmingensis sp. nov. differs from P. boettgeri by its smaller body size (vs SVL 34.5–37.8 mm in males and 39.7–46.8 mm in females), heels overlapping (vs just meeting), and round light patches on the shoulder absent (vs present). Panophrys sanmingensis sp. nov. differs from P. huangshanensis by the smaller body size (vs SVL 36.0– 41.6 mm in males and 44.2 mm in female), heels overlapping (vs just meeting or not meeting), wide lateral fringes and rudimentary web on toes present (vs both absent), and round light patches on the shoulder absent (vs present). Panophrys sanmingensis sp. nov. differs from P. kuatunensis by its heels overlapping (vs not meeting), lateral fringes on toes wide (vs narrow), and rudimentary web on toes present (vs absent). Panophrys sanmingensis sp. nov. differs from P. lishuiensis by the smaller body size (vs SVL 30.7–34.7 mm in males and 36.9–40.4 mm in female), tongue notched (vs not notched), wide lateral fringes and rudimentary web on toes present (vs both absent). Panophrys sanmingensis sp. nov. differs from P. ombrophila by the tongue notched (vs not notched), heels overlapping (vs not meeting), and wide lateral fringes and rudimentary web on toes present (vs both absent). Panophrys sanmingensis sp. nov. differs from P. xianjuensis by the smaller body size (vs SVL 31.0– 36.3 mm in males and 41.6 mm in female), tongue notched (vs not notched), and lateral fringes on toes wide (vs narrow). With a small body size, SVL 27.0– 29.5 mm in males and 29.5 mm in female, Panophrys sanmingensis sp. nov. is different from 34 congeners whose SVL> 32 mm in males or> 36 mm in females, namely P. angka, P. baolongensis, P. binchuanensis, P. binlingensis, P. brachykolos, P. caobangensis, P. caudoprocta, P. daweimontis, P. fansipanensis, P. hoanglienensis, P. insularis, P. jiangi, P. jingdongensis, P. jinggangensis, P. leishanensis, P. liboensis, P. lini, P. minor, P. mirabilis, P. mufumontana, P. nankunensis, P. obesa, P. omeimontis, P. palpebralespinosa, P. sangzhiensis, P. shuichengensis, P. shunhuangensis, P. spinata, P. tuberogranulatus, P. wugongensis, P. wuliangshanensis, P. wushanensis, P. xiangnanensis, and P. yangmingensis. Panophrys sanmingensis sp. nov. can be distinguished from the remaining seven congeners by the following characteristics: SVL 27.0– 29.5 mm in males and 29.5 mm in females (vs SVL 31.8–34.1 mm in females in P. cheni; SVL 30.2–39.3 mm in males in P. dongguanensis; vs SVL 30.4–33.9 mm in males and 34.1–37.5 in females in P. jiulianensis; vs SVL 30.5–37.3 mm in males in P. nanlingensis); horn-like tubercle at upper eyelid small (vs large in P. acuta); vomerine teeth absent (vs present in P. dongguanensis, P. jiulianensis, P. nanlingensis, P. rubrimera, and P. shimentaina); tongue notched (vs not notched in P. acuta, P. dongguanensis, and P. shimentaina); heels overlapping (vs not meeting in P. acuta and P. dongguanensis); wide lateral fringes present on toes (vs narrow in P. acuta, P. nanlingensis, P. rubrimera, and P. shimentaina; vs absent in P. dongguanensis and P. jiulianensis); rudimentary web present between toes (vs absent in P. rubrimera). Description of holotype. SYS a002498, adult male. Habitus small, SVL 29.3 mm; head length shorter than head width, HDW/HDL 1.02; snout rounded in dorsal view, projecting, sloping posteriorly to mouth in profile, protruding well beyond margin of lower jaw; dorsal surface of head flat; eye large, ED/HDL 0.42; nostril obliquely ovoid; pupil vertical; canthus rostralis well developed, curved above nostril; loreal region sloping; internasal distance larger than interorbital distance; tympanum distinct; choanae large ovoid, situated at base of maxilla; vomerine teeth absent; margin of tongue notched. Lower arm length 0.25 of SVL and hand length 0.24 of SVL; relative finger lengths I Coloration of holotype. Dorsal surface brown; a dark incomplete triangular marking with light edge between eyes; a dark “X” shaped marking with light edge on center of dorsum; dark crossbars on dorsal upper arms and hindlimbs; dark stripes below eyes and at lateral tip of snout; iris yellowish brown. Ventral surface pale with densely-distributed white spots; three dark longitudinal stripes on the throat; a pair of longitudinal black stripes with white edge on the lateroventral belly; palms and soles purplish brown, tips of digits greyish white, metacarpal and metatarsal tubercles orange red; pectoral glands and femoral glands white. Variation. Measurements of type series are given in Table 6. All specimens were similar in morphology. SYS a002493 (Fig. 5E) has reddish brown dorsal surface, with red spots on the dorsum and flanks. SYS a002494 (Fig. 5F) and SYS a002495 have black spots on flanks and the posterior of dorsum. Distribution and ecology. Currently, Panophrys sanmingensis sp. nov. is recognized from the Mt Longqi (700–740 m a.s.l.) of Jiangle County, Mt Emeifeng (900–1250 m a.s.l.) of Taining County and Gutian Town (750– 780 m a.s.l.) of Shanghang County in western Fujian, Mt Magu (500–600 m a.s.l.) of Nancheng County, Mt Junfeng (850–980 m a.s.l.) of Nanfeng County and Mt Jinpen (ca 500 m a.s.l.) of Xinfeng County in eastern Jiangxi, and Mt Fenghuang (800–900 m a.s.l.) of Chao’an District, Chaozhou City in eastern Guangdong, which indicates the distribution area of this species is in the wide hilly area among Fujian, Jiangxi and Guangdong provinces. This species inhabits streams surrounded by moist subtropical secondary evergreen broadleaved forests. Males call actively from mid-April to early June in Mt Longqi, Mt Emeifeng, Mt Magu, and Mt Junfeng; two individuals were found from Gutian Town in mid-July, and they were calling inactively; only one individual was found from Mt Jinpen in late September, and it was not calling., Published as part of Lyu, Zhi-Tong, Zeng, Zhao-Chi, Wang, Jian, Liu, Zu-Yao, Huang, Ya-Qiong, Li, Wen-Zhou & Wang, Ying-Yong, 2021, Four new species of Panophrys (Anura, Megophryidae) from eastern China, with discussion on the recognition of Panophrys as a distinct genus, pp. 9-40 in Zootaxa 4927 (1) on pages 28-31, DOI: 10.11646/zootaxa.4927.1.2, http://zenodo.org/record/4533849, {"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"]}

Panophrys daoji Lyu, Zeng, Wang & Wang sp. nov. Chresonymy. Megophrys sp20— Liu et al. 2018 Holotype. SYS a006209 (Fig. 4 A–D), adult male, collected by Jian Wang and Zhao-Chi Zeng on 31 July 2017 from Huading (29.2781°N, 121.0996°E; ca 680 m a.s.l.), Mt Tiantai, Tiantai County, Taizhou City, Zhejiang Province, PR China. Paratypes. Three adult males (SYS a006210–6211, SYS a006212/ CIB 116076) and two adult females (SYS a006213–6214), collected at the same time from the same locality as the holotype. Etymology. The specific epithet daoji is used as a noun in apposition, and refers to the Master Daoji (ḋằñ ®), also known as Ji Gong (ằẌ). He purportedly possessed supernatural powers which he used to help the poor and stand up to injustice, and became a famous legend in Chinese culture and a well-known deity in Chinese folk religion. Master Daoji was born in Yongning Village situated at the foot of Mt Tiantai where is the type locality of this new species with variable coloration. Common names. Daoji’s Horned Toad (in English) / Dào Jì Jiăo Chán (ḋằfflḃ in Chinese) Diagnosis. (1) body size moderate, with SVL 32.6–33.6 mm (n = 4) in adult males and 37.5–41.4 mm (n = 2) in adult females; (2) tympanum edge distinct and raised, upper margin in contact with supratympanic fold; (3) vomerine teeth absent; (4) margin of tongue not notched; (5) hindlimbs short, heels not meeting; (6) tibio-tarsal articulation reaching at center of tympanum; (7) TIB/SVL 0.39–0.42, FTL/SVL 0.57–0.64; (8) narrow lateral fringes on fingers present, one subarticular tubercle present at the base of each finger, relative finger lengths II Comparisons. Comparative data of Panophrys daoji sp. nov. with P. daiyunensis, and 47 recognized Panophrys congeners are given in Table 3. Panophrys daoji sp. nov. differs from P. daiyunensis by the relatively larger body size, SVL 32.6–33.6 mm in males and 37.5–41.4 mm in females (vs SVL 27.6–28.7 mm in males and 33.7–35.6 mm in females), vomerine teeth absent (vs present), heels not meeting (vs overlapping or meeting), and nuptial spines present on the finger I in breeding males (vs absent). Compared with the six Panophrys species previously recorded from eastern China, P. daoji sp. nov. differs from P. boettgeri by its smaller body size in adult males (vs SVL 34.5–37.8 mm), tongue not notched (vs notched), and round light patches on the shoulder absent (vs present). Panophrys daoji sp. nov. differs from P. huangshanensis by the smaller body size (vs SVL 36.0– 41.6 mm in males and 44.2 mm in female), tongue not notched (vs notched), narrow lateral fringes and rudimentary web on toes present (vs both absent), and round light patches on the shoulder absent (vs present). Panophrys daoji sp. nov. differs from P. kuatunensis by the larger body size in adult males (vs SVL 26.2–29.6 mm), tongue not notched (vs notched), and rudimentary web on toes present (vs absent). Panophrys daoji sp. nov. differs from P. lishuiensis by the narrow lateral fringes and rudimentary web on toes present (vs both absent). Panophrys daoji sp. nov. differs from P. ombrophila by its larger body size in adult females (vs SVL 32.8–35.0 mm), and narrow lateral fringes and rudimentary web on toes present (vs both absent). Panophrys daoji sp. nov. differs from P. xianjuensis by the heels not meeting (vs overlapping). With a moderate body size, SVL 32.6–33.6 mm in males and 37.5–41.4 mm in females, Panophrys daoji sp. nov. can be distinguished from 20 congeners whose SVL 35 mm in males or SVL 45 mm in females, namely P. acuta, P. baolongensis, P. binlingensis, P. caudoprocta, P. cheni, P. hoanglienensis, P. insularis, P. jingdongensis, P. jinggangensis, P. liboensis, P. mirabilis, P. obesa, P. omeimontis, P. palpebralespinosa, P. sangzhiensis, P. shuichengensis, P. spinata, P. tuberogranulatus, P. xiangnanensis, and P. yangmingensis. Panophrys daoji sp. nov. can be distinguished from the remaining 21 congeners by the following characteristics: vomerine teeth absent (vs present in P. daweimontis, P. dongguanensis, P. fansipanensis, P. jiulianensis, P. nankunensis, P. nanlingensis, P. rubrimera, and P. shimentaina); tongue not notched (vs notched in P. fansipanensis, P. jiulianensis, P. minor, P. nanlingensis, and P. rubrimera); heels not meeting (vs overlapping or meeting in P. angka, P. binchuanensis, P. jiangi, P. jiulianensis, P. leishanensis, P. lini, P. minor, P. mufumontana, P. nanlingensis, P. shimentaina, P. shunhuangensis, P. wuliangshanensis, and P. wushanensis); narrow lateral fringes present on toes (vs absent in P. angka, P. brachykolos, P. caobangensis, P. daweimontis, P. dongguanensis, P. fansipanensis, P. jiangi, P. jiulianensis, P. leishanensis, P. minor, P. nankunensis, P. shunhuangensis, P. wugongensis, and P. wuliangshanensis; vs wide in P. binchuanensis and P. lini; vs absent in females while wide in males in P. wushanensis); rudimentary web present between toes (vs lacking webs in P. daweimontis, P. fansipanensis, P. rubrimera, and P. wuliangshanensis). Description of holotype. SYS a006209, adult male. Habitus moderate, SVL 33.4 mm; head width shorter than head length, HDW/HDL 0.99; snout rounded in dorsal view, projecting, sloping posteriorly to mouth in profile, protruding well beyond margin of lower jaw; dorsal surface of head flat; eye small, ED/HDL 0.39; nostril obliquely ovoid; pupil vertical; canthus rostralis well developed, curved above nostril; loreal region sloping; internasal distance larger than interorbital distance; tympanum edge distinct and raised, upper margin in contact with supratympanic fold; choanae large ovoid, situated at base of maxilla; vomerine teeth absent; margin of tongue not notched. Lower arm length 0.20 of SVL and hand length 0.23 of SVL; relative finger lengths II Coloration of holotype. Dorsal surface dark brown; scarlet granules on the dorsal head, supratympanic fold, dorsum, and flanks; a dark brown triangular marking with light edge between eyes; a dark brown “X” shaped marking with light edge on center of dorsum; dark stripes below eyes; iris red. Ventral surface grayish brown, with a large white patch and tiny scarlet spots on the belly; three dark longitudinal stripes on the throat; palms pale, soles grayish brown, tips of digits greyish white, metacarpal and metatarsal tubercles pale; pectoral glands and femoral glands white. Variation. Measurements of type series are given in Table 5. All specimens were similar in morphology but their colorations were variable. Females are larger than males. SYS a006210 is dark gray on dorsum with “)(”- shaped skin ridge and “)(”-shaped marking on center of dorsum. SYS a006211 (Fig. 4E) is pale brown dorsally with underdeveloped “)(”-shaped skin ridge on center of dorsum. SYS a006212 is yellowish brown on dorsum. SYS a006214 (Fig. 4F) is reddish brown on dorsum with light patches on the shoulder. Distribution and ecology. Currently, Panophrys daoji sp. nov. is known from the Mt Tiantai (600–700 m a.s.l.) of Tiantai County and neighboring Xikou Town (500–600 m a.s.l.) of Fenghua City, both situated in the Tiantai Mountains in eastern Zhejiang. Males call from banks of streams in July, but tadpoles have not been found and additional ecological information remains unknown., Published as part of Lyu, Zhi-Tong, Zeng, Zhao-Chi, Wang, Jian, Liu, Zu-Yao, Huang, Ya-Qiong, Li, Wen-Zhou & Wang, Ying-Yong, 2021, Four new species of Panophrys (Anura, Megophryidae) from eastern China, with discussion on the recognition of Panophrys as a distinct genus, pp. 9-40 in Zootaxa 4927 (1) on pages 25-26, DOI: 10.11646/zootaxa.4927.1.2, http://zenodo.org/record/4533849, {"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"]}

Panophrys daiyunensis Lyu, Wang & Wang sp. nov. Chresonymy. Megophrys sp18— Liu et al. 2018 Holotype: SYS a001733 (Fig. 3 A–D), adult male, collected by Run-Lin Li on 22 May 2012 from Daiyun Village (25.6362°N, 118.2139°E; ca 1040 m a.s.l.), Daiyun Mountain Nature Reserve, Dehua County, Quanzhou City, Fujian Province, PR China. Paratypes: Three adult males (SYS a001730/ CIB 116075, SYS a001731–1732), collected at the same time from the same locality as the holotype; one adult female (SYS a006002), collected by Jian Wang on 26 June 2017 from the same locality as the holotype; two adult females (SYS a006000, 6003), collected by Zhi-Tong Lyu, Ying-Yong Wang, and Ya-Qiong Huang on 26 June 2017 from Jiuxianshan (25.7101°N, 118.1200°E; ca 1200 m a.s.l.), Daiyun Mountain Nature Reserve. Etymology. The specific epithet daiyunensis refers to the type locality of the new species, the Daiyun Mountain Nature Reserve. Common names. Daiyun Horned Toad (in English) / Dài Yún Jiǎo Chán (ăủfflḃ in Chinese) Diagnosis. (1) body size small, with SVL 27.6–28.7 mm (n = 4) in adult males and 33.7–35.6 mm (n = 3) in adult females; (2) tympanum edge distinct, upper margin in contact with supratympanic fold, TD/ED 0.45–0.53; (3) vomerine teeth present; (4) margin of tongue not notched; (5) heels overlapping or meeting; (6) tibio-tarsal articulation reaching just posterior to eye; (7) TIB/SVL 0.36–0.42, FTL/SVL 0.51–0.60; (8) narrow lateral fringes on fingers present, one subarticular tubercle present at the base of each finger, relative finger lengths I = II Comparisons. Comparative data of Panophrys daiyunensis sp. nov. with 47 recognized Panophrys congeners are given in Table 3. Six Panophrys species occur in eastern China, namely P. boettgeri, P. huangshanensis, P. kuatunensis, P. lishuiensis, P. ombrophila, and P. xianjuensis. Panophrys daiyunensis sp. nov. differs from P. boettgeri by the smaller body size, SVL 27.6–28.7 mm in males and 33.7–35.6 mm in females (vs SVL 34.5–37.8 mm in males and 39.7– 46.8 mm in females), vomerine teeth present (vs absent), tongue not notched (vs notched), and round light patches on the shoulder absent (vs present). Panophrys daiyunensis sp. nov. differs from P. huangshanensis by its smaller body size (vs SVL 36.0– 41.6 mm in males and 44.2 mm in female), vomerine teeth present (vs absent), tongue not notched (vs notched), narrow lateral fringes and rudimentary web on toes present (vs both absent), and round light patches on the shoulder absent (vs present). Panophrys daiyunensis sp. nov. differs from P. kuatunensis by its vomerine teeth present (vs absent), tongue not notched (vs notched), heels overlapping or meeting (vs not meeting), and rudimentary web on toes present (vs absent). Panophrys daiyunensis sp. nov. differs from P. lishuiensis by the smaller body size (vs SVL 30.7–34.7 mm in males and 36.9–40.4 mm in female), vomerine teeth present (vs absent), and narrow lateral fringes and rudimentary web on toes present (vs both absent). Panophrys daiyunensis sp. nov. differs from P. ombrophila by its vomerine teeth present (vs absent), heels overlapping or meeting (vs not meeting), and narrow lateral fringes and rudimentary web on toes present (vs both absent). Panophrys daiyunensis sp. nov. differs from P. xianjuensis by the smaller body size (vs SVL 31.0– 36.3 mm in males and 41.6 mm in female), and vomerine teeth present (vs absent). With a small body size, SVL 27.6–28.7 mm in adult males and 33.7–35.6 mm in adult females, Panophrys daiyunensis sp. nov. can be distinguished from 32 congeners whose SVL> 31 mm in males or> 38 mm in females, namely P. angka, P. baolongensis, P. binchuanensis, P. binlingensis, P. brachykolos, P. caobangensis, P. caudoprocta, P. daweimontis, P. fansipanensis, P. hoanglienensis, P. insularis, P. jiangi, P. jingdongensis, P. jinggangensis, P. leishanensis, P. liboensis, P. lini, P. mirabilis, P. minor, P. nankunensis, P. obesa, P. omeimontis, P. palpebralespinosa, P. sangzhiensis, P. shuichengensis, P. spinata, P. tuberogranulatus, P. wugongensis, P. wuliangshanensis, P. wushanensis, P. xiangnanensis, and P. yangmingensis. Panophrys daiyunensis sp. nov. can be distinguished from the remaining nine congeners by the following characteristics: SVL 27.6–28.7 mm in males and 33.7–35.6 mm in females (vs SVL 30.2–39.3 mm in males in P. dongguanensis; vs SVL 30.4–33.9 mm in males in P. jiulianensis; vs SVL 30.1–30.8 mm in males and 36.3 mm in female in P. mufumontana; vs SVL 30.5–37.3 mm in males in P. nanlingensis; vs SVL 30.3–33.7 mm in males and 37.6 mm in female in P. shunhuangensis); horn-like tubercle at upper eyelid small (vs large in P. acuta); vomerine teeth present (vs absent in P. acuta, P. cheni, P. mufumontana, and P. shunhuangensis); tongue not notched (vs notched in P. cheni, P. jiulianensis, P. nanlingensis, and P. rubrimera); heels overlapping or meeting (vs not meeting in P. acuta and P. dongguanensis); narrow lateral fringes present on toes (vs wide in P. cheni; vs absent in P. dongguanensis, P. jiulianensis, and P. shunhuangensis); rudimentary web present between toes (vs absent in P. rubrimera). Description of holotype. SYS a001733, adult male. Habitus small, SVL 28.7 mm; head width shorter than head length, HDW/HDL 0.97; snout rounded in dorsal view, projecting, sloping posteriorly to mouth in profile, protruding well beyond margin of lower jaw; dorsal surface of head flat; eye small, ED/HDL 0.39; nostril obliquely ovoid; pupil vertical; canthus rostralis well developed, curved above nostril; loreal region sloping; internasal distance larger than interorbital distance; tympanum distinct, small, upper margin in contact with supratympanic fold; choanae large ovoid, situated at base of maxilla; vomerine teeth present; margin of tongue not notched. Lower arm length 0.23 of SVL and hand length 0.24 of SVL; relative finger lengths I = II Coloration of holotype. Dorsal surface yellowish brown; a dark incomplete triangular marking with light edge between eyes; a dark “X” shaped marking with light edge on center of dorsum; dark patches on dorsal upper arms and hindlimbs; dark stripes below eyes and at lateral tip of snout; iris reddish brown. Ventral surface dark brown; three dark longitudinal stripes on the throat, middle one distinctly shorter; red spots on the chest and densely-distributed tiny white spots on belly and ventral thigh; palms and soles purplish brown, tips of digits greyish white, metacarpal and metatarsal tubercles orange red; pectoral glands and femoral glands white. Variation. Measurements of type series are given in Table 4. All specimens were similar in morphology. Females are larger than males. SYS a006000 (Fig. 3E, F) has reddish brown dorsal surface with “)(”-shaped marking and gray ventral surface with unclear marking. Without “X” or “)(”-shaped marking on center of dorsum in SYS a006002. Distribution and ecology. Currently, Panophrys daiyunensis sp. nov. is known from Daiyun Mountain Nature Reserve (1000–1250 m a.s.l.) and Xiamen City (ca 400 m a.s.l.) of southern Fujian. This toad inhabits streams surrounded by moist subtropical secondary evergreen broadleaved forests, and is common from May to June. Males call actively on leaves of bushes or rocks near streams during this period. All females found in June were gravid with oocytes but tadpoles have not been found., Published as part of Lyu, Zhi-Tong, Zeng, Zhao-Chi, Wang, Jian, Liu, Zu-Yao, Huang, Ya-Qiong, Li, Wen-Zhou & Wang, Ying-Yong, 2021, Four new species of Panophrys (Anura, Megophryidae) from eastern China, with discussion on the recognition of Panophrys as a distinct genus, pp. 9-40 in Zootaxa 4927 (1) on pages 21-23, DOI: 10.11646/zootaxa.4927.1.2, http://zenodo.org/record/4533849, {"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"]}

Panophrys tongboensis Wang & Lyu sp. nov. Chresonymy. Megophrys sp21— Liu et al. 2018 Holotype. SYS a003228 (Fig. 6 A–E), adult male, collected by Run-Lin Li on 18 August 2014 from Pingxi (28.1154°N, 118.2372°E; ca 1100 m a.s.l.), Mt Tongbo, Guangfeng District, Shangrao City, Jiangxi Province, PR China. Paratypes. Three adult males (SYS a003225–3227), collected at the same time from the same locality as the holotype; one adult male (SYS a001911/ CIB 116078), collected by Run-Lin Li on 5 August 2012 from the same locality as the holotype. Etymology. The specific epithet tongboensis refers to its type locality, Mt Tongbo. Common names. Mt Tongbo Horned Toad (in English) / Tóng Bó Shān Jiǎo Chán (ffiůƜfflḃin Chinese) Diagnosis. (1) body size small, with SVL 26.5–31.5 mm (n = 5) in adult males; (2) tympanum visible, the anterior edge indistinct; (3) vomerine teeth present; (4) margin of tongue notched; (5) heels overlapping; (6) tibiotarsal articulation reaching at center of eye; (7) TIB/SVL 0.46–0.49, FTL/SVL 0.59–0.68; (8) narrow lateral fringes on fingers present, one subarticular tubercle present at the base of each finger, relative finger lengths II Comparisons. Comparative data of Panophrys tongboensis sp. nov. with P. daiyunensis, P. daoji, P. sanmingensis, and 47 recognized Panophrys congeners are given in Table 3. Panophrys tongboensis sp. nov. differs from P. daiyunensis by its tongue notched (vs not notched), and lateral fringes and web on toes absent (vs both present). Panophrys tongboensis sp. nov. differs from P. daoji by the smaller body size, SVL 26.5–31.5 mm in males (vs SVL 32.6–33.6 mm), vomerine teeth present (vs absent), tongue notched (vs not notched), heels overlapping (vs not meeting), and lateral fringes and web on toes absent (vs both present). Panophrys tongboensis sp. nov. differs from P. sanmingensis by its vomerine teeth present (vs absent), and lateral fringes and web on toes absent (vs both present). Compared with the six Panophrys species previously recorded from eastern China, P. tongboensis sp. nov. differs from P. boettgeri by its smaller body size (vs SVL 34.5–37.8 mm in males), vomerine teeth present (vs absent), heels overlapping (vs just meeting), lateral fringes and web on toes absent (vs both present), and round light patches on the shoulder absent (vs present). Panophrys tongboensis sp. nov. differs from P. huangshanensis by the smaller body size (vs SVL 36.0– 41.6 mm in males), vomerine teeth present (vs absent), heels overlapping (vs just meeting or not meeting), and round light patches on the shoulder absent (vs present). Panophrys tongboensis sp. nov. differs from P. kuatunensis by vomerine teeth present (vs absent), heels overlapping (vs not meeting), and lateral fringes on toes absent (vs present). Panophrys tongboensis sp. nov. differs from P. lishuiensis by its smaller body size (vs SVL 30.7–34.7 mm in males), vomerine teeth present (vs absent), and tongue notched (vs not notched). Panophrys tongboensis sp. nov. differs from P. ombrophila by its vomerine teeth present (vs absent), tongue notched (vs not notched), and heels overlapping (vs not meeting). Panophrys tongboensis sp. nov. differs from P. xianjuensis by its vomerine teeth present (vs absent), tongue notched (vs not notched), and lateral fringes and web on toes absent (vs both present). With a small body size, SVL 26.5–31.5 mm in males, Panophrys tongboensis sp. nov. is different from 21 congeners whose SVL> 34 mm in males, namely P. baolongensis, P. binlingensis, P. caobangensis, P. caudoprocta, P. daweimontis, P. hoanglienensis, P. insularis, P. jiangi, P. jingdongensis, P. jinggangensis, P. liboensis, P. lini, P. minor, P. mirabilis, P. obesa, P. omeimontis, P. palpebralespinosa, P. sangzhiensis, P. shuichengensis, P. spinata, and P. xiangnanensis. Panophrys tongboensis sp. nov. can be distinguished from the remaining 20 congeners by the following characteristics: SVL 26.5–31.5 mm in males (vs SVL 32.0–36.0 mm in P. binchuanensis; vs SVL 33.2–37.1 mm in P. yangmingensis; vs SVL 33.2–39.0 mm in P. tuberogranulatus; vs SVL 33.7–39.3 mm in P. brachykolos); horn-like tubercle at upper eyelid small (vs large in P. acuta); vomerine teeth present (vs absent in P. acuta, P. angka, P. binchuanensis, P. brachykolos, P. cheni, P. leishanensis, P. mufumontana, P. shunhuangensis, P. tuberogranulatus, P. wugongensis, P. wuliangshanensis, P. wushanensis, and P. yangmingensis); tongue notched (vs not notched in P. acuta, P. angka, P. brachykolos, P. dongguanensis, P. leishanensis, P. mufumontana, P. nankunensis, P. shimentaina, P. shunhuangensis, P. tuberogranulatus, P. wugongensis, P. wushanensis, and P. yangmingensis); heels overlapping (vs not meeting in P. acuta, P. brachykolos, P. dongguanensis, P. nankunensis, and P. wugongensis); lateral fringes on toes absent (vs present in P. acuta, P. binchuanensis, P. cheni, P. mufumontana, P. nanlingensis, P. rubrimera, P. shimentaina, and P. yangmingensis; vs absent in females while wide in males in P. wushanensis); web between toes absent (vs present in P. acuta, P. angka, P. binchuanensis, P. brachykolos, P. cheni, P. dongguanensis, P. jiulianensis, P. leishanensis, P. mufumontana, P. nankunensis, P. nanlingensis, P. shimentaina, P. shunhuangensis, P. tuberogranulatus, P. wugongensis, P. wushanensis, and P. yangmingensis); one subarticular tubercle present at the base of each finger (vs absent in P. fansipanensis). Description of holotype. SYS a003228, adult male. Habitus small, SVL 28.5 mm; head width shorter than head length, HDW/HDL 0.91; snout rounded in dorsal view, projecting, sloping posteriorly to mouth in profile, protruding well beyond margin of lower jaw; dorsal surface of head flat; eye large, ED/HDL 0.39; nostril obliquely ovoid; pupil vertical; canthus rostralis well developed, curved above nostril; loreal region sloping; internasal distance larger than interorbital distance; tympanum visible but the anterior edge indistinct; choanae large ovoid, situated at base of maxilla; vomerine teeth present; margin of tongue notched. Lower arm length 0.24 of SVL and hand length 0.24 of SVL; relative finger lengths II Shank length 0.46 of SVL and foot 0.61 of SVL; tibio-tarsal articulation reaches at center of eye when hindlimb stretched alongside body; heels overlapping when hindlimbs held at right angles to body; relative toe lengths I Dorsal body’s skin texture relatively smooth with tiny granules; “)(”-shaped skin ridge on center of dorsum; a small horn-like tubercle present at edge of upper eyelid; supratympanic fold distinct, curving from posterior corner of eye, posteroventrally to above insertion of arm; flank with raised tubercles. Ventral surface smooth, with scattered tubercles on thigh; pectoral gland large, closer to axilla; single large femoral gland on posterior surface of thigh. Coloration of holotype. Dorsal surface beige; a brown “V” shaped marking with light edge between eyes; a brown “)(”-shaped marking with light edge on center of dorsum; dark crossbars on dorsal upper arms and hindlimbs; dark stripes below eyes and at lateral tip of snout; iris reddish brown. Ventral surface dark brown with densely-distributed white spots; a dark longitudinal stripe on the throat; a pair of longitudinal black stripes with white edge on the lateroventral belly; palms and soles reddish brown, tips of digits light red, metacarpal and metatarsal tubercles orange red; pectoral glands and femoral glands white. Variation. Measurements of type series are given in Table 7. All specimens were similar in morphology. SYS a003227 has an incomplete triangular marking between eyes. SYS a001911 (Fig. 6F) has a triangular marking between eyes and short longitudinal skin ridges on the flanks. Distribution and ecology. Currently, Panophrys tongboensis sp. nov. is only known from the type locality, Mt Tongbo (1100–1115 m a.s.l.) in northeastern Jiangxi. This toad appears to be rare with all individuals being found in the same stream, and is under the competition with the sympatric congener P. boettgeri which is more abundant. Males call on deadwood above streams in August, however, the male individuals found in August were not bearing nuptial pads or spines. Females and tadpoles have not been found and additional ecological information remains unknown.

The diversity of Panophrys horned toads is considered highly underestimated with a large number of undescribed cryptic species. In this work, we describe four Panophrys species from eastern China which were proposed as cryptic species by molecular data in previous study, additionally provide new information on the biogeography of these four species. Panophrys daiyunensis sp. nov. from southern Fujian, Panophrys daoji sp. nov. from eastern Zhejiang, Panophrys sanmingensis sp. nov. from the hilly area among Fujian, Jiangxi and Guangdong, and Panophrys tongboensis sp. nov. from northeastern Jiangxi, can be distinguished from all recognized congeners by a combination of morphological characteristics. The descriptions of these four new species take the recognized species of Panophrys to 51, which is the largest genus within the Asian horned toads subfamily Megophryinae. Considered as an appropriate arrangement for the Asian horned toads currently and applied in this study to describe the new species, the generic recognition of Panophrys is also discussed.

Nidirana occidentalis sp. nov. Lyu, Yang, and Wang Chresonymy Rana pleuraden— Yang and Rao 2008 (Longling, Baoshan, and Tengchong) Pelophylax pleuraden— Fei et al. 2009 (Longling, Baoshan, and Tengchong) Dianrana pleuraden— Fei et al. 2012 (Western Yunnan) Babina pleuraden— Chan and Bi 2016 (Tengchong) Nidirana pleuraden— Lyu et al. 2017 (Mt. Gaoligong) Holotype. SYS a003776, adult male, collected by Jian Wang and Hai-Long He on 15 May 2015 from Dahaoping (24.9846°N, 98.7376°E; ca 1600 m a.s.l.), Mt. Gaoligong, Yunnan Province, PR China. Paratypes. Males SYS a003775/ CIB 116075, SYS a003777 and female SYS a003778, collected at the same time from the same locality as the holotype. Males SYS a007830–7831 and Females SYS a007829, 7832, collected by Jian Wang, Yu-Long Li and Yao Li on 9 June 2019 from Mengku Township (23.6586°N, 99.9847°E; ca 1980 m a.s.l.), Shuangjiang Lahu, Va, Blang and Dai Autonomous County, Yunnan Province, PR China. Etymology. The specific name “ occidentalis ” is a Latin adjective and means “western”, referring to the distribution of the new species in western Yunnan, which takes the westernmost distribution among all congeners. Common name. Western Yunnan Music Frog (in English) / Dian Xi Qin Wa (AEOiḪü in Chinese) Diagnosis. Nidirana occidentalis sp. nov. is placed in the genus Nidirana based on the morphological characteristics of the absence of the thumb-like structure on finger I, presence of well-developed dorsolateral folds, and the presence of suprabrachial glands in breeding males (Lyu et al. 2017), and is further assigned to the N. pleuraden group by the absence of lateroventral groove on all digits (Dubois 1992; Lyu et al. 2019). Nidirana occidentalis sp. nov. is distinguished from all known Nidirana congeners by the following combination of the morphological characteristics: (1) body medium-size and elongated, SVL 49.1 ± 3.3 (44.5–53.0, N = 5) mm in adult males and 58.8 ± 2.9 (55.6–61.3, N = 3) mm in adult females; (2) head relatively short, HDW/HWL 0.84 ± 0.05 in males and 0.86 ± 0.04 in females; (3) tympanum relatively small, TD/ED 0.81 ± 0.02 in males and 0.77 ± 0.02 in females (4) lateroventral grooves absent on every digit; (5) absence of supernumerary tubercles on hand; (6) tibio-tarsal articulation reaching at the eye; (7) heels just meeting; (8) posterior of dorsal skin rough with tubercles and spinules; (9) a pair of subgular vocal sacs in males; (10) one single nuptial pad present on the first finger in breeding males, nuptial spinules invisible; (11) suprabrachial gland large and smooth; (12) calling: 3–5 identical regular notes. Description of holotype: SYS a003776 (Fig. 6), adult male. Body elongated, SVL 47.5 mm; head relatively short, flat above; snout rounded in dorsal and lateral views, slightly protruding beyond lower jaw, longer than horizontal diameter of eye (SNT/ED 1.33); canthus rostralis distinct, loreal region concave; nostril round, closer to the snout than to the eye; a longitudinal swollen mandibular ridge extending from below nostril through lower edges, eye and tympanum to above insertion of arm; supratympanic fold absent; interorbital space flat, narrower than internasal distance (IND/IOD 1.29); pupil elliptical, horizontal; tympanum distinct, round, relatively small, and close to eye; pineal ocellus invisible; vomerine ridge present, bearing small teeth; tongue large, cordiform, notched behind. Presence of a pair of subgular vocal sacs, a pair of slit-like openings at posterior of jaw. Forelimbs moderately robust, lower arm 17% of SVL and hand 28% of SVL; fingers thin, relative finger lengths II Hindlimbs relatively robust, tibia 51% of SVL and foot 82% of SVL; heels just meeting when hindlimbs flexed at right angles to axis of body; tibio-tarsal articulation reaching at the eye when hindlimb is stretched along the side of the body; toes relatively long and thin, relative lengths I Dorsal skin of head and anterior body smooth, posterior dorsum of body rough with dense tubercles and horny spinules; developed dorsolateral fold from posterior margin of upper eyelid to above groin but intermittent posteriorly; flank smooth; a large and smooth suprabrachial gland behind base of forelimb, not prominent; dorsal surfaces of thigh and tibia with several longitudinal ridges and bearing spinules. Ventral surface of head, body, and limbs smooth; large flattened tubercles densely arranged on the rear of thigh and around vent. Coloration of holotype. Photos of the holotype in life were not taken and the coloration data is lacking. In preservative, dorsal surface of head and body dark brown; a longitudinal light mid-dorsal stripe extending from snout to vent; dorsolateral fold bicolor, upper part light brown and lower part black; flank light brown with irregular black spots; suprabrachial gland creamy. Dorsal forelimbs light brown; a longitudinal black stripe on the anterior surface of the forelimb; irregular black marks on dorsal surface of the forelimb; dorsal hindlimbs reddish brown with black crossbars on the thigh, tibia and tarsus; nuptial pad grey. Loreal and temporal regions dark brown; tympanum light brown. Lips and throat white; ventral surface of body and limbs creamy white; rear thigh tinged with pink; ventral hand and foot pale. Variations. Measurements of type series are given in Table 3 and photos of paratypes are shown in Fig. 7. All type series specimens were similar in morphology. Females are significantly larger than males, with the sexual size dimorphism of 83.5%. SYS a007830–7832 have several tubercles on the upper flank, not bearing spinules. Suprabrachial glands prominent in SYS a007830 (vs. not prominent in holotype SYS a003776 and paratype SYS a007831). The coloration of Nidirana occidentalis sp. nov. is variable, from yellowish brown, reddish brown to dark brown on dorsum; pineal ocellus invisible; a light mid-dorsal stripe edged with broad darker stripes extending from forehead to vent; black spots scatter on posterior dorsum. Dorsolateral fold bicolor, light color upper and darker lower part; flank varies from light brown to dark brown, with small black spots or large black patches. One black stripe in front of the base of forelimb; irregular dark marks on lateral forelimb; three to four dark brown or black crossbars on thigh, tibia and tarsus. Temporal region lighter color with a yellowish arc patch in SYS a007830, 7831; maxillary gland and shoulder gland white; upper ⅓ iris brownish white and lower ⅔ iris reddish brown. Ventral surface of body and limbs creamy white; several black spots on the edge of ventral belly and limbs in SYS a007832. Surface of throat darker; ventral hand and foot purplish brown. Distribution and ecology. Currently, Nidirana occidentalis sp. nov. is recognized from multiple localities in western Yunnan, all of which are situated to the west of the Red River (Fig. 1). This frog inhabits natural or artificial ponds surrounded by moist subtropical secondary evergreen broadleaved forests, and is common in its habitat, sympatric with Tylototriton shanjing Nussbaum, Brodie, and Yang, 1995, Nanorana yunnanensis (Anderson, 1879), and Zhangixalus puerensis (He, 1999). The adult males were heard calling in the water surface from May to July.

Two new toad species of the genus Leptobrachella are described from the Yunnan-Guizhou Plateau of China, based on the combination of molecular and morphological data. The description of Leptobrachella aspera Wang, Lyu, Qi & Wang, sp. nov. from Huanglianshan Nature Reserve represents the thirteenth Leptobrachella species known from Yunnan Province, and the description of Leptobrachella dorsospina Wang, Lyu, Qi & Wang, sp. nov. from Yushe Forest Park represents the sixth Leptobrachella species known from Guizhou Province. These new discoveries further emphasize the extremely high diversity of the Leptobrachella toads in these regions.

Two new toad species of the genusLeptobrachellaare described from the Yunnan-Guizhou Plateau of China, based on the combination of molecular and morphological data. The description ofLeptobrachella asperaWang, Lyu, Qi & Wang,sp. nov.from Huanglianshan Nature Reserve represents the thirteenthLeptobrachellaspecies known from Yunnan Province, and the description ofLeptobrachella dorsospinaWang, Lyu, Qi & Wang,sp. nov.from Yushe Forest Park represents the sixthLeptobrachellaspecies known from Guizhou Province. These new discoveries further emphasize the extremely high diversity of theLeptobrachellatoads in these regions.

Recent phylogenetic analysis encompassing multilocus nuclear-gene and matrilineal mtDNA genealogy has revealed a series of cryptic species of the subgenus Panophrys within genus Megophrys from southern and eastern China. This study demonstrates that the Panophrys specimens from the hilly areas among Guangdong, Guangxi and Hunan can be morphologically distinguished from all recognized congeners, thereby providing additional supports for the recognitions of four new species of Panophrys, namely Megophrys (Panophrys) mirabilis Lyu, Wang & Zhao, sp. nov. from northeastern Guangxi, Megophrys (Panophrys) shimentaina Lyu, Liu & Wang, sp. nov. from northern Guangdong, and Megophrys (Panophrys) xiangnanensis Lyu, Zeng & Wang, sp. nov. and Megophrys (Panophrys) yangmingensis Lyu, Zeng & Wang, sp. nov. from southern Hunan. The descriptions of these species take the number of Megophrys species to 101, 46 of which belong to the subgenus Panophrys.

Amolops shuichengicus sp. nov., a new species of the A. mantzorum group is described from Guizhou, southwest China, on the basis of significant molecular divergences in 16S + CO1 genes and the combination of morphological characteristics: small body size, SVL 34.6–39.6 mm in adult males and 48.5–55.5 mm in adult females; dorsal skin relatively smooth; presence of vomerine teeth; presence of cream maxillary gland from lower edge of eye to the anterior of supratympanic fold; presence of supratympanic folds and glandular dorsolateral folds; tympanum indistinct; absence of a circummarginal groove on the disk of the first finger; presence of supernumerary tubercles below the base of fingers III and IV; absence of outer metatarsal tubercle and tarsal glands; males without vocal sacs. In addition, evidenced by the phylogenetic analyses in this study and literature data, we suggest that A. liangshanensis should be synonymized with A. loloensis and the records of A. marmoratus in Yunnan, China should be referred to A. afghanus. Following our proposal, the genus Amolops contains 57 species, with 32 recorded from China.

Amolops yunkaiensis Lyu, Wang, Liu, Zeng and Wang sp. nov. (Fig. 3 and 4) Chresonymy: Amolops torrentis (Smith, 1923): Fei et al. 2009 (Xinyi, Guangdong). Amolops torrentis: Wei et al. 2010 (Yunkaishan Nature Reserve, Xinyi, Guangdong). Holotype: SYS a004705, adult male, collected by Zhi-Tong Lyu (ZTL), Jian Wang (JW), and Hai-Long He (HLH) on 20 April 2016 from Bajiaxianhu (21��53���35.5���N, 111��29���5.1���E; 441 m a.s.l.), Ehuangzhang Nature Reserve, Guangdong Province, China. Paratypes: 16 adult specimens. Male SYS a000773 and female SYS a000774, collected on 29 March 2010 by Run-Lin Li from the same stream as holotype; six females, SYS a004698���4701, 4706���4707, collected on 19���20 April 2016 by ZTL, JW and HLH from the same stream as holotype. Two females SYS a003979 and 3982 collected on 14 May 2015 by ZTL, Chao Huang and Kai-Chi Huang from Liping (2148���59.9���N, 11126���57.5���E; 337 m a.s.l.), EHZ, Guangdong Province, China. Six males, SYS a004674, 4681���4684, collected on 15���16 April 2016 by ZTL, Ying-Yong Wang and HLH, and SYS a004992/ CIB 106880 collected on 29 June 2016 by JW and HLH, from Dongkeng (2214���50.3���N, 11111���23.1���E; 987 m a.s.l.), Yunkaishan Nature Reserve, Guangdong Province, China. Etymology. The specific name, yunkaiensis, refers to the locality of the new species, the Yunkai Mountains. We suggest its English common name ���Yunkai Torrent Frog��� and Chinese name ���Yun Kai Tuan Wa (ĀH�����)���. Diagnosis. The diagnostic character for the genus Amolops is the presence of an abdominal sucker in the tadpole (Rao & Wilkinson 2007). However, because the tadpole of the new species remains to be discovered, we assigned the new species to this genus based on the morphological and genetic similarity of the adult specimens to those of A. ricketti and A. wuyiensis. Amolops yunkaiensis sp. nov. is distinguished from its congeners by a combination of the following morphological characteristics: (1) relatively small body size, SVL 31.8���34.1 mm in adult males, 35.2���39.0 mm in adult females; (2) dorsal skin of body very rough with numerous raised large warts; (3) several large raised warts on flanks, white or yellowish white; (4) dorsal body olive-brown or light brown with dark brown blotches; (5) vomerine teeth absent; (6) tongue cordiform, deeply notched posteriorly; (7) absence of the dorsolateral folds; (8) presence of a circummarginal groove on the disk of the first finger; (9) absence of outer metatarsal tubercles; (10) absence of tarsal glands; (11) sparse translucent tubercles on the lower jaw, forechest, posterior of belly and ventral thigh in males; (12) males with a pair of subgular vocal sacs; (13) nuptial pad on first finger prominent with developed white conical spines in breeding males. Description of Holotype: Head length slightly larger than head width (HDW/HDL = 0.93); snout short (SNT/ HDL = 0.36) and rounded in profile, projecting beyond lower jaw; nostril closer to tip of snout than eye; loreal region concave; top of head flat; eye large (ED/HDL = 0.4) and convex; canthus rostralis distinct; pineal body barely visible; tympanum small, edge faintly distinct; supratympanic fold distinct, from back of eye to shoulder; choanae moderate; vomerine teeth absent; tongue cordiform, deeply notched posteriorly. Forelimbs moderately robust; hands moderately long (HND/SVL = 0.31); relative finger lengths I Measurement of holotype (in mm). SVL 32.4; HDL 12.3; HDW 11.5; SNT 4.4; IND 3.9; IOD 2.8; ED 4.9; TD 1.5; TED 1.3; HND 10.1; RAD 6.3; FTL 23.2; TIB 17.5; F3W 2.3; T4W 2.1. Color in life. Dorsal body olive-brown with dark brown blotches; large warts on flanks yellowish white; a dark brown longitudinal band on each side of lower flank of body; faint dark transverse bars on dorsal surface of limbs; dorsal discs of digits greyish-white; posterior edge of upper lip and rictal gland light maize-yellow. Throat, chest, and belly creamy white; tubercles on the lower jaw, forechest, posterior of belly and ventral thigh slightly translucent; ventral surfaces of limbs with greyish-white flecks; creamy white blotches on ventral thighs; rear of thighs mottled with dark brown; ventral hands and feet dark grey; a yellowish oval spot on each side of cloaca. Color in preservative. Dorsal surface of head, body and limbs transformed into dark olive with black blotches, transverse bars and spots; ventral surface of head and body white ventral surface of limbs light brown. Variation. Measurements of type series are given in Table 4. All paratypes are very similar in morphology and color pattern except that dorsal skin yellowish brown in SYS a003979 and 4674; throat or chest with grey flecks in SYS a000774, 3983, 3984, 4699, 4703, 4704 and 4707; pineal body distinctly visible in SYS a000773, 0 776, 0 778, 0 779, 3979, 4670, 4681, 4683, 4704, 4706 and 4992; heel just meeting in SYS a004700, 4701 and 4993; tibiotarsal articulation reaching tip of snout when hindlimb stretched alongside of body in SYS a000773, 0 778, 3983, 3984, 4674, 4681, 4682 and 4683. Male secondary sexual characteristics. A pair of subgular vocal sacs, a pair of slit-like openings at posterior of jaw; a white nuptial pad on first finger prominent with strongly developed white conical spines. Comparisons. Amolops yunkaiensis sp. nov. was previously reported as A. torrentis, but differs from true A. torrentis from Hainan Island by having numerous raised large warts on dorsum and flanks (vs. smooth in A. torrentis), tarsal glands absent (vs. present), having sparse translucent tubercles on the lower jaw, forechest, posterior of belly and ventral thigh in males (vs. absent), having nuptial pad prominent with developed conical spines on first finger in males (vs. nuptial pad and spines absent). Amolops yunkaiensis sp. nov. differs from A. albispinus, A. ricketti, A. wuyiensis, A. daiyunensis, A. hongkongensis and A. hainanensis, to which it is most closely related (Fig. 2), by having a relatively small body size, SVL 31.8���34.1 mm in adult males and SVL 35.2���39.0 mm in adult females (vs. SVL 36.7���42.4 mm in adult males and SVL 43.1���51.9 mm in adult females of A. albispinus, SVL 42.0��� 60.5 mm in adult males and SVL 53.5��� 67.0 mm in adult females of A. ricketti, SVL 38.0���45.0 mm in adult males and SVL 45.0���53.0 mm in adult females of A. wuyiensis, SVL 34.0���41.0 mm in adult males and SVL 31.0���48.0 mm in adult females of A. hongkongensis, SVL 36.0���58.0 mm in adult males and SVL 44.0���63.0 mm in adult females of A. daiyunensis, SVL 71.0���93.0 mm in adult males and SVL 68.0���78.0 mm in adult females in A. hainanensis); the presence of numerous raised large warts on dorsal body in A. yunkaiensis (vs. absent in A. ricketti, A. wuyiensis and A. daiyunensis); tarsal glands absent in A. yunkaiensis (vs. present in A. daiyunensis and A. hainanensis); having translucent tubercles on the lower jaw, forechest, posterior of belly and ventral thigh in males (vs. absent in A. albispinus, A. ricketti, A. daiyunensis, A. hongkongensis and A. hainanensis); nuptial pad on first finger prominent with developed white conical spines (vs. absent in A. hainanensis, black conical spines in A. wuyiensis); having a pair of vocal sacs in A. yunkaiensis (vs. absent in A. albispinus, A. ricketti and A. hainanensis). Amolops yunkaiensis sp. nov. differs from the remaining congeners in having tubercles on the lower jaw, forechest, posterior of belly and ventral thigh in males; the absence of dorsolateral folds (vs. present in A. akhaorum, A. aniqiaoensis, A. archotaphus, A. bellulus, A. chakrataensis, A. chayuensis, A. chunganensis, A. compotrix, A. cremnobatus, A. cucae, A. gerbillus, A. iriodes, A. jaunsari, A. kohimaensis, A. longimanus, A. mengyangensis, A. minutus, A. monticola, A. nyingchiensis, and A. vitreus); the presence of a circummarginal groove on disk of first finger (vs. absent in A. formosus, A. granulosus, A. jinjiangensis, A. xinduqiao, A. liangshanensis, A. lifanensis, A. loloensis, A. mantzorum, A. nidorbellus, A. tuberodepressus, and A. viridimaculatus); the absence of vomerine teeth (vs. present in A. afghanus, A. assamensis, A. caelumnoctis, A. daorum, A. himalayanus, A. indoburmanensis, A. kaulbacki, A. larutensis, A. marmoratus, A panhai, A. spinapectoralis and A. splendissimus); having dorsal surface olive-brown with dark brown blotches (vs. dorsal color of green or bright yellow in A. medogensis). Distribution and ecology. Amolops yunkaiensis sp. nov. inhabits rocky, fast-flowing streams (300���900 m a.s.l.) surrounded by moist subtropical secondary evergreen broadleaved forests and the frog is sympatric with a congener, A. ricketti, in the same steams. All individuals were observed from April to June, when males were found bearing mature nuptial pads. Nevertheless, the call of the frog was not heard during any of our repeated surveys, and much of its ecology and behavior remains unknown. Currently, the Yunkai Torrent Frog is only known from the type locality, the Ehuangzhang Nature Reserve and Yunkaishan Nature Reserve in the Yunkai Mountains. Its extent of occurrence is estimated to be less than 5000 km 2, and the area of occupancy is estimated to be less than 500 km 2. Moreover, these areas are being threatened by hydropower station construction and tourism development. This frog appeared to be rare in EHZ and YKS. Surveys are urgently needed in southern China to investigate the population status and the distribution of this species. We suggest the species may be eligible for listing as Endangered (B1ab(i, ii, iii) + 2ab(i, ii, iii)) in the IUCN Red List of Threatened Species., Published as part of Lyu, Zhi-Tong, Wu, Jun, Wang, Jian, Sung, Yik-Hei, Liu, Zu-Yao, Zeng, Zhao-Chi, Wang, Xin, Li, You-Yu & Wang, Ying-Yong, 2018, A new species of Amolops (Anura: Ranidae) from southwestern Guangdong, China, pp. 562-576 in Zootaxa 4418 (6) on pages 569-573, DOI: 10.11646/zootaxa.4418.6.4, http://zenodo.org/record/1245414, {"references":["Smith, M. A. (1923) On a collection of reptiles and batrachians from the island of Hainan. Journal of the Natural History Society of Siam, 6 (2), 195 - 212.","Fei, L., Hu, S. Q., Ye, C. Y. & Huang, Y. Z. (2009) Fauna Sinica. Amphibia Vol. 2 Anura. Science Press, Beijing, 957 pp.","Wei, Q. L., He, B. & Hu, H. J. (2010) Amolops torrentis, a new record of Amphian from Guangdong. Sichuan Journal of Zoology, 29 (3), 345.","Rao, D. Q. & Wilkinson, J. A. (2007) A new species of Amolops (Anura: Ranidae) from southwest China. Copeia, 2007 (4), 913 - 919. https: // doi. org / 10.1643 / 0045 - 8511 (2007) 7 [913: ansoaa] 2.0. co; 2"]}

Gracixalus guangdongensis sp. nov. Fig. 3 Holotype. SYS a005724, adult male, collected on 13 April 2017 by Jian Wang (JW), Zhao-Chi Zeng (ZCZ) and Dian-Cheng Yang (DCY) from DWL (22��17���31.27��� N, 111��12���50.42���E; 1600 m a.s.l.) in Maoming City, Guangdong Province, China. Paratypes. Two adult females were collected from the same locality as the holotype: SYS a004686, collected by Ying-Yong Wang (YYW), Zhi-Tong Lyu (ZTL), JW and ZCZ on 16 April 2016; SYS a004698, collected by JW, ZCZ, Can-Rong Lin (CRL) and Chun-Peng Guo (CPG) on 28 June 2016. Eight adult males: SYS a004672���4673, SYS a004685, 4687���4689, collected by Hai-Long He (HLH), Run-Lin Li (RLL) and ZTL on 16 April 2016 from the same locality as the holotype; SYS a004983/ CIB106882, collected by JW, ZTL and ZCZ on 28 June 2016, and SYS a005241, collected by JW on 16 August 2016 from DWL (22��15���57.58��� N, 111��11���18.10��� E; 1000 m a.s.l.). Three adult males: SYS a004902���4904, collected by YYW and ZTL on 5 June 2016 from MNK (23��38���16.40��� N, 113��50���49.40��� E; 600 m a.s.l.) in Huizhou City, Guangdong Province, China. Five adult males: SYS a005750��� 5751, collected by YYW and ZTL on 20 April 2017 and SYS a005773���5775, collected by JW, ZTL and HLH on 5 May 2017, all from NNR (24��55���53.82��� N, 113��01���19.66��� E; 1000 m a.s.l.) in Shaoguan City, Guangdong Province, China. Referred specimen. A single juvenile of the species (SYS a004671), collected at the same locality as the holotype by ZCZ on 15 April 2016. Etymology. The specific epithet ��� guangdongensis ��� refers to the distribution of this species, Guangdong Province, China. We propose the common English name ��� Guangdong Tree Frog���, Chinese name ���Guang Dong Xian Shu Wa (���������������)��� for this species. Diagnosis. Gracixalus guangdongensis sp. nov. is assigned to genus Gracixalus based upon our preliminary mtDNA phylogenetic analyses and the following morphological characters: the presence of an intercalary cartilage between the terminal and penultimate phalanges of digits, tips of digits expanded into large discs bearing circum marginal grooves, vomerine teeth absent, horizontal pupil, tibia length greater than four times width, distance between nostrils less than or equal to between eyes, rictal gland connected to the mouth (Delorme et al. 2005; Rowley et al. 2014), and the back bearing dark X-or inverted V-shape (Fei et al. 2009). Gracixalus guangdongensis sp. nov. can be distinguished from its congeners by a combination of the following morphological characters: (1) relatively small body size (SVL 26.1���34.7 mm in adult males, 34.9���35.4 mm in adult females); (2) upper eyelid and dorsum lacking spines; (3) supratympanic fold and tympanum distinct; (4) dorsal and lateral surface rough, sparsely scattered with tubercles; (5) ventral skin granular; (6) tibiotarsal projection absent; (7) toe-webbing moderately developed, finger webbing rudimentary; (8) heels slightly overlapping when flexed hindlimbs are held at right angles to the body axis; (9) brown to beige above, with an inverse Y-shaped dark brown marking extendeing from the interorbital region to the centre of the dorsum; (10) males with a single subgular vocal sac and protruding nuptial pads with minute granules on the dorsal surface of the base of first finger. Description of holotype. Adult male, dorsoventrally compressed; head width almost equal to head length (HDW:HDL 1.01); snout triangularly pointed in dorsal view, projecting beyond margin of the lower jaw; canthus rostralis distinct and rounded; loreal region oblique and concave; nostrils oval, significantly protuberant, closer to tip of snout than to the eye; interorbital region flat, interorbital distance slightly broader than internasal distance (IOD: IND 1.03); eye large, horizontal diameter slightly smaller than snout length (ED:SL 0.83); pupil horizontal; tympanum distinct, tympanic rim weakly developed; tympanum diameter smaller than half of eye diameter (TYD:ED 0.41); supratympanic fold distinct, extending from the posterior corner of the eye above insertion of arm; vomerine teeth absent; tongue deeply notched behind; a single subgular vocal sac present. Forelimb moderately robust; forearm and hand relatively long (FAHL:SVL 0.48), hand significantly longer than forearm (HAL:SVL 0.46); fingers dorsoventrally compressed, webbing rudimentary, with narrow lateral fringes on outer edge of all fingers; relative finger length I Measurements of holotype (in mm). SVL 30.2, HDL 10.7, HDW 10.8, SL 5.1, IND 3.5, IOD 3.6, ED 4.2, TYD 1.7, TED 0.8, HAL 9.8, FAHL 14.6, TIBL 14.1, FTL 20.7, FDW III 2.1, FPW III 1.0, TDW IV 1.9, TPW IV 1.2. Coloration of holotype in life. Dorsal surface brown, with an inverse Y-shaped, dark-brown marking, starting at the interorbital region, bifurcating into two branches on shoulder, extending posteriorly; dorsal surface of fore- and hindlimbs with dark brown transverse bands; several faint, small, black blotches scattered on ventrolateral region of flanks and on ventral surface; ventral surface of hindlimbs, throat and chest creamy white, surface of belly and forelimbs light brown; iris brown, pupil black. Coloration of holotype in preservative. Color of dorsal surface dark grey; ventral surface of head and body greyish white; the Y-shaped marking and transverse black bands, blotches on the ventrolateral region of the flanks and the ventral surface are greyish black, and more distinct than in life. Variation. The external morphology and color pattern of all paratypes corresponds to that of the holotype, except for the following characters: background color of dorsal surface of head, body and limbs is beige in SYS a004671 (the single juvenile), SYS a004672 and SYS a004686 (vs. brown in holotype SYS a005724); adult males collected in the breeding season (SYS a004672���4673, 4685, 4687���4689, 4983, 4902���4904, 5724, 5750���5751, 5773���5775) possess nuptial pads on the dorsal surface of first finger, but the single specimen collected in the nonbreading season (SYS a005241) has barely visible nuptial pads; specimens from MNK and NNR have a smaller body size than specimens from DWL, measurement and body proportions of the type series specimens of G. guangdongensis sp. nov. are given in Table 3. Comparisons. Gracixalus guangdongensis sp. nov. was assigned with strong node supporting values (0.82 in BI and 86% in ML) to Clade I in our phylogenetic tree containing G. jinxiuensis, G. jinggangensis, G. sapaensis, G. nonggangensis and G. waza. The new species differs from G. jinxiuensis by the relatively larger body size, SVL 26.1���34.7 mm in adult males, 34.9���35.4 mm in adult females (vs. 24.2���26.3 mm in males, 28���29.2 mm in females of G. jinxiuensis), more developed webbing (formula of webbing of the fourth toes in (3) IV (3)) (vs. (3��) IV (3��) in G. jinxiuensis), ventral surface of hand, foot and palm smooth with sparse small tubercles (vs. rough with dense large tubercles in G. jinxiuensis), tibio-tarsal articulation reaching the middle of the eye (vs. reaching the posterior corner of eye the in G. jinxiuensis), and having heels overlapping when the flexed hindlimbs are held at right angles to the body axis (vs. just meeting in G. jinxiuensis); from G. jinggangensis by the relatively larger body size in females, SVL 34.9���35.4 mm (vs. 31.6 mm in G. jinggangensis), presence of nuptial pad on the first finger of male (vs. on first and second fingers of male G. jinggangensis), and heels overlapping when the flexed hindlimbs are held at right angles to the body axis (vs. just meeting in G. jinggangensis); from G. sapaensis by the presence of a distinct tympanum (vs. indistinct in G. sapaensis), presence of a single vocal sac (vs. paired vocal sac in in G. sapaensis), brown to beige background color of the dorsal surface of head, body and limbs in life (vs. greyish to reddish brown in G. sapaensis), creamy white to light brown ventral surface (vs. venter pink in G. sapaensis), and rough dorsal surface (vs. nearly smooth in G. sapaensis); from G. nonggangensis by the more developed webbing (formula of webbing of the fourth toes in (3) IV (3)) (vs. (3��) IV (3��) in G. nonggangensis), ventral surface of hand, foot and palm smooth with sparse small tubercles (vs. rough with dense large tubercles in G. nonggangensis), presence of nuptial pad on the first finger of males (vs. absent in G. nonggangensis), presence of rudimentary finger webbing (vs. finger webbing absent in G. nonggangensis), tibio-tarsal articulation reaching the middle of the eye (vs. reaching the tip of the snout in G. nonggangensis), and brown to beige background color of the dorsal surface of the head, body and limbs in life (vs. yellowish olive in G. nonggangensis); from G. waza by the relatively small body size in adult females, SVL 34.9���35.4 mm (vs. 37.6 mm in G. waza), brown to beige background color of the dorsal surface of the head, body and limbs in life (vs. greyish green to moss-green in G. waza), and rough dorsal skin with tubercles (vs. nearly smooth dorsal skin in G. waza). Gracixalus guangdongensis sp. nov. can also be easily distinguished from the six members of Clade II (Fig 2). It differs from G. seesom by the relatively larger body size, SVL 26.1���34.7 mm in adult males, 34.9���35.4 mm in adult females (vs. 21.6���23 mm in males and 23.2���25.4 mm in female G. seesom), presence of nuptial pads on the first finger of males (vs. absence in male G. seesom), and rough dorsal skin with tubercles (vs. nearly smooth dorsal skin in G. seesom); from G. quangi by the relatively larger body size, SVL 26.1���34.7 mm in adult males, 34.9���35.4 mm in adult females (vs. 21.4���22.9 mm in males and 26.8���27.3 mm in females of G. quangi), absence of spines on the upper eyelid (vs. presence in G. quangi), absence of a tibiotarsal projection (vs. presence in G. quangi), and brown to beige background color of the dorsal surface of the head, body and limbs in life (vs. greenish to brownish green in G. quangi); from G. gracilipes by the brown to beige background color of dorsal surface of head, body and limbs and the presence of inverse Y-shaped dark brown marking on back in life (vs. transparent green with Xshaped brown markings in G. gracilipes), absence of a white patch under the eye to the tympanum (vs. presence in G. gracilipes), and absence of spines on upper eyelid (vs. presence in G. gracilipes); from G. supercornutus by the brown to beige background color of the dorsal surface (vs. yellowish green in G. supercornutus), the absence of a white patch extending from under the eye to the tympanum (present in G. supercornutus), absence of spines on upper eyelid (vs. present in G. supercornutus), absence of a tibiotarsal projection (vs. present in G. supercornutus); from G. quyeti by having distinct supratympanic fold (vs. indistinct in G. quyeti), brown to beige background color of dorsal surface of head, body and limbs (vs. brown to moss-green in G. quyeti); from G. lumarius by the relatively smaller body size, SVL 26.1���34.7 mm in adult males, 34.9���35.4 mm in adult females (vs. 38.9���41.6 mm in males and 36.3 mm in female of G. lumarius), distinct tympanum and supratympanic fold (vs. indistinct in G. lumarius), skin on dorsal surface of head, body rough, sparsely scattered with brown tubercles (vs. dorsum with distinctive dense network of white conical tubercles in adult males in G. lumarius), venter creamy white to greyish white (vs. venter pink in G. lumarius), single vocal sac in males (vs. a pair of vocal sacs in G. lumarius). The new species also differs from the two congeners for which comparative molecular material is not available. The new species differs from Gracixalus megdogensis by the presence of rudimentary finger webbing (vs. absence of finger webbing in G. megdogensis), tibio-tarsal articulation reaching the middle of the eye (vs. reaching the anterior corner of the eye in G. megdogensis), rough dorsal skin with tubercles (vs. smooth dorsal skin in G. megdogensis), and brown to beige background color of dorsal surface of head, body and limbs in life (vs. green in G. megdogensis); from G. carinensis by the relatively smaller body size in females, SVL 34.9���35.4 mm (vs. 38 mm in G. carinensis), more developed webbing (webbing formula of the fourth toe (3) IV (3)) (vs. webbing formula (2��) IV (2) in G. carinensis), and snout length slightly longer than diameter of eye (SL> ED) (vs. snout length shorter than diameter of eye (SL G. carinensis). Gracixalus guangdongensis sp. nov. further differs from Kurixalus ananjevae (Matsui & Orlov), which is morphologically and likely molecularly similar to the Gracixalus species of Clade II (i.e. Clade I in this study) according to Rowley et al. 2011, by the smaller body size in females, SVL 34.9���35.4 mm (vs. 43.0 mm in female K. ananjevae), dark grey dorsum in preservation (vs. pinkish gray in preservation in K. ananjevae), snout length slightly longer than diameter of eye (SL> ED) (vs. snout length equal to or shorter than diameter of eye (SL ��� ED) in K. ananjevae), and more developed webbing (webbing formula of the fourth toes is (3) IV (3)) (vs. (3) IV (2��) in K. ananjevae). Gracixalus guangdongensis sp. nov. further differs from the small- and medium-sized rhacophorids occurred in southern China and northern Vietnam: the new species differs from Liuixalus calcarius Milto, Poyarkov, Orlov & Nguyen, L. feii Yang, Rao & Wang, L. jinxiuensis Li, Mo, Jiang, Xie & Jiang, L. hainanus (Liu and Wu), L. ocellatus (Liu & Hu), L. romeri (Smith), L. shiwandashan Li, Mo, Jiang, Xie & Jiang, Raorchestes longchuanensis (Yang and Li) and Ra. menglaensis (Kou) by the significantly larger body size, SVL 26.1���34.7 mm in adult males, 34.9���35.4 mm in adult females (vs. SVL values not-overlaping and less than 21.0 mm in males and 22.0 mm in females) and the more developed webbing on toes (vs. very poorly-developed webbing on the toes); from Philautus abditus Inger, Orlov & Darevsky by the presence of nuptial pads in males and a distinct tympanum (vs. nuptial pads absent and tympanum completely hidden under skin); from Buergeria oxycephala (Boulenger) by the significantly smaller body size and the absence of vomerine teeth (vs. SVL 34���38 in males and 60���68 in females, vomerine teeth present); from Thelerdema albopunctatum (Liu & Hu), T. annae Nguyen, Pham, Nguyen, Ngo & Ziegler, T. bicolor (Bourret), T. corticale (Boulenger), T. gordoni Taylor, T. lateriticum Bain, Nguyen & Doan and T. rhododiscus (Liu &Hu) by the absence of skin ridges and large warts on dorum (vs. presence of irregular skin ridges and large warts on dorsum in T. bicolor, T. corticale, T. gordoni; and presence of irregular skin ridges on dorsum in T. albopunctatum and T. rhododiscus), presence of a single vocal sac (vs. paired vocal sac in T. albopunctatum, and absent in T. annae, T. lateriticum and T. rhododiscus), and the absence of vomerine teeth (vs. vomerine teeth present in T. bicolor and T. corticale); from Kurixalus bisacculus (Taylor) by the presence of a single vocal sac in males and fingers with rudiment of web (vs. paired vocal sac in males and fingers without web), from K. lenquanensis Yu, Wang, Hou, Rao & Yang and K. odontotarsus (Ye and Fei) by the absence of vomerine teeth and absence of conical tubercles on posterolateral surface of forearms and tibia (vs. vomerine teeth present, distinct conical tubercles on posterolateral surface of forearms and tibia present); from Chiromantis doriae (Boulenger) by having rough dorsal skin with tubercles and a ���Y-shaped��� marking on the back (vs. smooth dorsal skin with granules and five longitudinal stripes on the back); from Feihyla fuhua Fei, Ye & Jiang, F. palpebralis (Smith) and F. vittata (Boulenger) by the single vocal sac in males (vs. paired vocal sac in males); from Polypedates braueri (Vogt), Po. impresus Yang, Po. megacephalus Hallowell and Po. mutus (Smith) by the absence of vomerine teeth and presence of rudiment of web on fingers (vs. vomerine teeth present and webbing on fingers absent); from Rhacophorus dorsoviridis Bourret, Rh. hoanglienensis Orlov, Lathrop, Murphy & Ho, Rh. laoshan Mo, Jiang, Xie & Ohler, Rh. larissae Ostroshabov, Orlov & Nguyen, Rh. leucofasciatus Liu & Hu, Rh. minimus Rao, Wilkinson & Liu, Rh. nigropunctatus Liu, Hu & Yan, Rh. pinglongensis, Published as part of Wang, Jian, Zeng, Zhao-Chi, Lyu, Zhi-Tong, Liu, Zu-Yao & Wang, Ying-Yong, 2018, Description of a new species of Gracixalus (Amphibia: Anura: Rhacophoridae) from Guangdong Province, southeastern China, pp. 251-269 in Zootaxa 4420 (2) on pages 257-265, DOI: 10.11646/zootaxa.4420.2.7, http://zenodo.org/record/1250885, {"references":["Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2005) Une nouvelle classification generique et subgenerique de la tribu des Philautini (Amphibia, Anura, Rhacophorinae). Bulletin Mensuel de la Societe Linneenne de Lyon, 74, 165 - 171. https: // doi. org / 10.3406 / linly. 2005.13595","Rowley, J. J. L., Le, D. T. T., Dau, V. Q., Hoang, H. D. & Cao, T. T. (2014) A striking new species of phytotelm-breeding tree frog (Anura: Rhacophoridae) from central Vietnam. Zootaxa, 3785 (1), 25 - 37. https: // doi. org / 10.11646 / zootaxa. 3785.1.2","Fei, L., Hu, S. Q., Ye, C. Y. & Huang, Y. Z. (2009) Fauna Sinica, Amphibia. Vol. 2. Anura. Science Press, Beijing, 957 pp.","Rowley, J. J. L., Dau, Q. V., Nguyen, T. T., Cao, T. T. & Nguyen, S. N. (2011) A new species of Gracixalus (Anura: Rhacophoridae) with a hyperextended vocal repertoire from Vietnam. Zootaxa, 3125, 22 - 38."]}

The diversity of the subgenus Panophrys within the genus Megophrys has been revealed to be extremely underestimated from southeastern China. Herpetological surveys coupled with extensive sampling in a longitudinal mountain belt located in southeastern China resulted in the discoveries of six new species of the subgenus Panophrys. Furthermore, the new discoveries support the findings of “micro-endemism”, “sympatric phenomenon” and “sympatric but distant phylogenetically” which appear to be common among Panophrys species, and also indicates that the Asian horned toads would be good candidates for studies on speciation and biogeography, and additionally emphasizes the conservation difficulties of these toads.

A new species of tree frog, G. guangdongensis sp nov., is described based on a series of specimens collected from Dawuling Forest Station, Mount Nankun and Nanling Nature Reserve of Guangdong Province, southeastern China. The new species is distinguished from all known congeners by a significant genetic divergence at the mitochondrial 16S rRNA gene fragment examined (p-distance ≥ 4.6%) and the following combination of morphological characters: relatively small body size (SVL 26.1–34.7 mm in adult males, 34.9–35.4 mm in adult females); upper eyelid and dorsum lacking spines; supratympanic fold and tympanum distinct; dorsal and lateral surface rough, sparsely scattered with tubercles; ventral skin granular; tibiotarsal projection absent; toe-webbing moderately developed, finger webbing rudimentary; heels slightly overlapping when flexed hindlimbs are held at right angles to the body axis; brown to beige above, with an inverse Y-shaped dark brown marking extendeing from the interorbital region to the centre of the dorsum; males with a single subgular vocal sac and protruding nuptial pads with minute granules on the dorsal surface of the base of first finger. The discovery and description of Gracixalus guangdongensis sp. nov. represents the 14th species known in this genus.

A new species, Amolops yunkaiensis sp. nov. is described based on a series of specimens from Ehuangzhang Nature Reserve and Yunkaishan Nature Reserve, southwestern Guangdong Province, China. The new species can be distinguished from all known congeners by molecular divergence in the mitochondrial 16S rRNA, 12S rRNA and CO1 genes, and a combination of the following characters: relatively small body size, SVL 31.8–34.1 mm in adult males, 35.2–39.0 mm in adult females; numerous raised large warts on dorsum and flanks; dorsal body olive-brown or light brown with dark brown blotches; absence of vomerine teeth; absence of tarsal glands; presence of a pair of subgular vocal sacs, nuptial spines on the first finger, and sparse translucent tubercles on the lower jaw, forechest, posterior belly and ventral thigh in male. Hence, the genus Amolops contains 52 species, 29 of which occur in China.

The Asian leaf litter toads of the genusLeptolalaxrepresent a highly diverse species group and currently contain 53 recognized species. During herpetological surveys in Yingjiang County, western Yunnan of China, we collected series ofLeptolalaxspecimens from an isolated small fragment of montane evergreen forest. Subsequent study based on acoustic, morphological and molecular data reveals that there were three different species among the specimens sampled: while one of them belongs toLeptolalax ventripunctataus, the other two species represent unknown taxa and are described herein:Leptolalax purpurussp. nov.andLeptolalax yingjiangensissp. nov. The two new species can be distinguished from other congeners by the molecular divergences, acoustic data, and by a combination of morphological characters including: body size, dorsal and ventral patterns, dorsal skin texture, sizes of pectoral and femoral glands, degree of webbing and fringing on the toes and fingers, dorsum coloration and iris coloration in life. Our results further reveal that species diversity of the genusLeptolalaxstill remains highly underestimated and warrants further attention.