Generic classification of Asian horned toads (Anura: Megophryidae: Megophryinae) and monograph of Chinese species.

The subfamily Megophryinae, as a representative batrachian group of the Oriental Realm and one of the most diverse groups of amphibians, has attracted considerable attention due to continued conjecture regarding its generic classification and failure to reach a satisfactory consensus. China boasts the richest diversity of Asian horned toads, containing some two thirds of the total species cataloged. However, most species have a complicated taxonomic history, resulting in multiple misidentifications. As such, an overall clarification of historical records and regional checklists is required. In the current investigation, we established the phylogeny of the Asian horned toads and performed detailed examinations with redefinitions of several important morphological traits. Based on the phylogenetic relationships and morphological differences, we propose a new ten-genus classification for the Asian horned toad subfamily Megophryinae: i.e., Brachytarsophrys, Atympanophrys, Grillitschia, Sarawakiphrys gen. nov., Jingophrys gen. nov., Xenophrys, Megophrys, Pelobatrachus, Ophryophryne, and Boulenophrys. Revisions on the diagnosability, distribution, and content of each genus are provided. Furthermore, we present a careful review of the taxonomic history of Asian horned toad species from China and provide a monograph of congeners, including six species of Brachytarsophrys, four species of Atympanophrys, five species of Jingophrys gen. nov., 10 species of Xenophrys, two species of Ophryophryne, and 60 species of Boulenophrys. Finally, we discuss the importance of traditional morphological traits based on multiple populations in taxonomic work as well as taxonomic inflation caused by the genetic species delimitation.

A new species of the genus Micryletta (Anura, Microhylidae) from Hainan Island, China.

We report on a new amphibian species of the genus Micryletta from Hainan Island, China, based on morphological and molecular analyses. The new species, Micryletta immaculata sp. nov., is diagnosed by a combination of the following morphological characters: medium-sized within genus (SVL 23.3-24.8 mm in males, n=3; 27.7-30.1 mm in females, n=2); dorsum bronze brown to reddish brown in life; dark brown spots and stripes on dorsum and flank absent; flanks largely pigmented with silver white; throat in adult males dark brown; undersides without dark patterns; outer metatarsal tubercle absent; supratympanic fold distinct; webbing between toes basal and poorly developed; tibiotarsal articulation adpressed limb reaching level of tympanum. The new species is divergent from all other congeners based on 16S rRNA gene sequences (3.0%-7.7%). Data on the natural history and male advertisement calls of the new species are provided. Following the IUCN Red List Categories and Criteria, we propose the new species to be listed as Vulnerable B1ab (iii).

Acoustic characteristics of the island-endemic Parapelophryne scalpta (Anura: Bufonidae) from Hainan Island, China.

The Hainan Pygmy Toad Parapelophryne scalpta (Liu Hu) is the single species of the genus Parapelophryne Fei, Ye Jiang in the Family Bufonidae (Frost 2020) and is endemic to China's Hainan Island, which lies within the tropics at 18°09'-20°10'N, 108°37'-111°03'E, with a land area of ca. 33,900 km2. The taxonomy status of this species has obtained much debates since its discovery: it was originally described by Liu et al. (1973) as a member of the genus Nectophryne Buchholz Peters then allocated to Pelophryne Barbour by Ye Fei (1978). On the basis of its unique morphological characters, Fei et al. (2003) erected a new genus Parapelophryne to discriminate P. scalpta from other bufonids. Some researchers, however, still listed the species under the genus Pelophryne (Stuart et al. 2008; Shi et al. 2011). Recently, the validity of the genus Parapelophryne is further confirmed by Matsui et al. (2015) on the basis of molecular analyses. Interestingly enough, Matsui et al. (2015)'s result showed that this extremely small-sized toad is a sister taxon of the large-sized toads of the genus Bufo Garsault, which is mainly distributed in temperate and subtropical regions of Eurasia (Frost 2020). Parapelophryne scalpta is a forest-dependent species and only inhabits in well-preserved evergreen broadleaf forests (Fei Ye 2016). It is the smallest toad species in China with snout-vent length of 19-23 mm in adult males and 24-27 mm in females (Fei Ye 2016). Due to their small size and elusive habits, little is known about the natural history of P. scalpta and its acoustic characteristics are still undocumented (Fei Ye 2016). During a herpetological survey in Hainan in 2015, we luckily detected male calls of P. scalpta and obtained a short but clear record of the advertisement call of the species. Herein, we describe the acoustic characteristics and calling behavior of Parapelophryne scalpta.

A new species of Nidirana from the N. pleuraden group (Anura, Ranidae) from western Yunnan, China.

The Music Frog genus Nidirana was recently resurrected as a distinct genus and contains 14 species distributed in subtropical eastern and southeastern Asia. The species diversity of Nidirana is dramatically underestimated, and half of its species was described in the last five years. In this study, Nidirana occidentalis sp. nov., a new species of Music Frog from western Yunnan, China, is proposed based on morphological and molecular evidences. The new species was previously misidentified as N. pleuraden, but can be distinguished from the true N. pleuraden from eastern Yunnan, and all other congeners, by a combination of morphological characteristics, and significant divergence in the mitochondrial genes (≥ 5.1% in 16S and ≥ 8.9% in CO1). Nidirana occidentalis sp. nov. is assigned to the N. pleuraden group on the basis of morphological characters, but its phylogenetic placement remains unresolved due to weak branch support. Geographically, these two species are isolated by the Red River in Yunnan, supporting the hypothesis that the Red River is an important geographical barrier that drives speciation in flora and fauna. Nidirana occidentalis sp. nov. represents the second species of N. pleuraden group and the 15th species of the genus.

Re-examination of the Chinese record of Opisthotropis maculosa (Squamata, Natricidae), resulting in the first national record of O. haihaensis and description of a new species.

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 hungtai sp. 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.

A new species of Amolops (Anura: Ranidae) from China, with taxonomic comments on A. liangshanensis and Chinese populations of A. marmoratus.

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.

A new limestone-dwelling species of Micryletta (Amphibia: Anura: Microhylidae) from northern Vietnam.

We report on a new species of the genus Micryletta from limestone karst areas in northern Vietnam, which is described on the basis of molecular and morphological evidence. Micryletta nigromaculata sp. nov. is restricted to narrow areas of subtropical forests covering karst massifs in Cat Ba National Park (Hai Phong Province) and Cuc Phuong National Park (Ninh Binh Province) at elevations of 90-150 m a.s.l. In the phylogenetic analyses, the new species is unambiguously positioned as a sister lineage to all remaining species of Micryletta. We also discuss genealogical relationships and taxonomic problems within the genus Micryletta, provide molecular evidence for the validity of M. erythropoda and discuss the taxonomic status of M. steinegeri. We suggest the new species should be considered as Endangered (B1ab(iii), EN) following the IUCN's Red List categories. A discussion on herpetofaunal diversity and conservation in threatened limestone karst massifs in Southeast Asia is provided.

Description of two new sympatric species of the genus Leptolalax (Anura: Megophryidae) from western Yunnan of China.

The Asian leaf litter toads of the genus Leptolalax represent a highly diverse species group and currently contain 53 recognized species. During herpetological surveys in Yingjiang County, western Yunnan of China, we collected series of Leptolalax specimens 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 to Leptolalax ventripunctataus, the other two species represent unknown taxa and are described herein: Leptolalax purpurussp. nov. and Leptolalax 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 genus Leptolalax still remains highly underestimated and warrants further attention.

A new species of the genus Megophrys (Anura: Megophryidae) from Yunnan Province, China.

We describe a new horned toad species of the genus Megophrys from Yingjiang County of Yunnan Province, China. The new species, Megophrys feii sp. nov., can be distinguished from its congeners by a combination of the following characters: (1) body slender and small (SVL 24.3-25.1 mm in four males, 28.2-28.9 mm in two females); (2) head length about equal to head width (HDL/HDW 0.97-0.99); (3) tympanum circular and distinct; (4) maxillary teeth present; (5) vomerine ridges and vomerine teeth absent; (6) tongue round, slightly notched posteriorly; (7) tibia long (TIB/SVL 0.48-0.52 in four males, 0.54-0.55 in two females), shanks overlapping when thighs are held at right angles to the body; (8) dorsal skin distinctly granular and densely covered with small tubercles; (9) lower flanks and lateral sides of belly scattered with small but prominent, white, tubercles; (10) webbing between toes rudimentary; (11) lateral fringes on toes moderate to wide; (12) breeding males without nuptial pads and spines on fingers; (13) protruding projection posterior to cloaca present in both sexes; (14) groin and ventral thigh coloration in life not contrasting with surrounding regions; (15) the presence of an indistinct and small horn-like tubercle at the edge of the eyelid; and (16) an advertisement call with a dominant frequency of 4.74-4.91 kHz (at 18 ˚C). To date, the new species has only been found at its type locality in stream areas in montane evergreen broadleaf forests between 700-1200 m elevation.

A new species of Hemiphyllodactylus (Squamata: Gekkonidae) from Hong Kong.

A new species of gekkonid, Hemiphyllodactylus hongkongensis sp. nov., is described based on a series of specimens collected from Hong Kong SAR, China. The new species can be distinguished from other known congeners by molecular divergence in the mitochondrial ND2 gene and morphological characters including 5-6 chin scales; a manual lamellar formula of 3-3(4)-4-4; a pedal lamellar formula of 3(4)-4(5)-4(5)-4; 24-25 continuous femoral and precloacal pores; 12-15 dorsal scales and 9-10 ventral scales contained in diameter of eye. At present, the genus Hemiphyllodactylus comprises 22 described species.

A new phytotelm-breeding treefrog of the genus Nasutixalus (Rhacophoridae) from western Yunnan of China.

A new treefrog of the genus Nasutixalus is described from montane evergreen forest in Yingjiang County, in western Yunnan, China. The new species, Nasutixalus yingjiangensis sp. nov., can be distinguished from its congeners by a combination of the following characters: medium body size (SVL 39.5-40.0 mm in adult males, 47.5 mm in a single female); tympanum indistinct and covered with tubercles; disc diameter of third finger greater than tympanum diameter; dorsal skin relatively smooth, scattered with small tubercles, those on head and anterior dorsum of body more dense and more prominent; light brown above with a dark brown marking between eyes and two broad dark brown lateral strips on the dorsum; iris with a weak "X"-shaped, light colored marking; interorbital distance shorter than the upper eyelid width; comparatively short foot (mean TFL/SVL ratio 67.0% and 62.9% in males and female respectively). The new species appears to be forest-dependent and was found in well-preserved montane rainforest; it inhabits the canopy and breeds in tree holes during the rainy season. The type locality of the new species is an isolated forest fragment surrounded by degraded landscape and efforts are already underway to protect the site. Nasutixalus yingjiangensis sp. nov. represents the third known species of the newly established genus Nasutixalus.

A new species of the genus Leptobrachium (Anura: Megophryidae) from the Gaoligongshan Mountain Range, China.

We describe a new species of the genus Leptobrachium from the Gaoligongshan Mountain Range, Yunnan Province of China based on molecular and morphological evidences. The new species, Leptobrachium tengchongense sp. nov., can be distinguished from its congeners by a combination of the following characters: (1) relatively small size (adult males SVL 41.7-51.5 mm); (2) head width slightly larger than head length; (3) tympanum indistinct; (4) two palmar tubercles oval and distinct, inner one larger than outer one; (5) sexually active males without spines on the upper lip; (6) dorsal skin smooth with distinct network of ridges; (7) dorsum pinkish grey and scattered with irregular black markings; (8) venter dark purplish-gray with numerous small white spots on tubercles, solid white chest; (9) iris bicolored, upper one-third light blue, lower two-third dark brown. With the description of the new species, the number of Leptobrachium species currently known from China adds up to ten.

A new species of the genus Leptolalax (Anura: Megophryidae) from Mt. Gaoligongshan of western Yunnan Province, China.

A new species of the genus Leptolalax is described from the Tengchong Section of Gaoligongshan National Nature Reserve, Tengchong County, Yunnan Province, China. The new species, Leptolalax tengchongensis sp. nov., can be distinguished from its congeners by a combination of the following characters: (1) small size (SVL 23.9-26.0 mm in males, 28.8-28.9 mm in females); (2) dorsal skin shagreened and scattered with fine, round reddish tubercles; (3) toes with rudimentary webbing and narrow lateral fringes; (4) tympanum distinctly discernible, almost entirely black; (5) ventrolateral glands indistinct; (6) flanks with several distinct and large dark blotches; (7) ventral surfaces white, scattered with distinct irregular dark speckling; (8) iris not bicolored, uniformly dark brown and scattered with minute, coppery reticulations throughout. To date, the new species has only been found at its type locality in evergreen broadleaf forests at elevations between 2000-2100 m.

A new species of the genus Odorrana (Amphibia: Ranidae) and the first record of Odorrana bacboensis from China.

The genus Odorrana currently contains at least 56 recognized species that inhabits montane streams in subtropical and tropical Asia. Twenty new species have been described in the last decade, indicating the potential cryptic species diversity of this genus. We collected several specimens of Odorrana species from Southern China from 2007 to 2014, and on the basis of a combined morphological characters and phylogenetic analysis, we described the new species Odorrana fengkaiensis sp. nov. herein. The new species is very similar to O. hainanensis and O. bacboensis, but can be consistently separated by morphology, and allopatric distribution. It is further reciprocally monophyletic to O. hainanensis in a mitochondrial gene trees with an average genetic divergence of 2.1% (1.9%-2.4%). The new species inhabits in lowland broad streams, rivers, pools and near the riparian areas, but its general ecology remains poorly known. The new species is characterized by its body length of adult females approximately twice as long as adult males (SVL 77.8-111.9 mm in females, 37.4-51.8 mm in males); eye large in males, eye diameter 1.01-1.16 times as long as snout length; tympanum of males large and distinct, extremely close to the eye, 0.7-1.4 mm in tympanum-eye distance; dorsolateral folds absent; dorsal skin shagreened, with several large tubercles in males; flanks with tubercles and scattered larger pustules, 8-10 of which usually arranged in a dorsolateral row; ventral skin smooth, with spines in adult males during the breeding season; the tibio-tarsal articulation stretched forward beyond the tip of snout; relative finger lengths: II < I < IV < III; dorsum brown with irregularly reticulated green markings in males and young females, uniformly brown in some old adult females; males with velvety nuptial pad on thumb, paired gular pouches; mature oocytes almost purely black in life, showed dark grey animal pole and olive vegetative pole in preservative. In addition, we found O. bacboensis, a new country record from China, indicating a range extension from north-central Vietnam to southeast Yunnan and adjacent area in Guangxi.

Two new species of the genus Goniurosaurus (Squamata: Sauria: Eublepharidae) from southern China.

Two new species of large geckos in the genus Goniurosaurus are described based on specimens collected from karst areas of Guangxi Zhuang Autonomous Region, southern China: Goniurosaurus kadoorieorum sp. nov. and Goniurosaurus kwangsiensis sp. nov. Data on natural history of the new species are provided, as well as discussions on the current conservation status of Goniurosaurus species in southern China. Due to the popularity of this genus as novelty pets, and recurring cases of scientific descriptions driving herpetofauna to near-extinction by commercial collectors, we do not disclose the collecting localities of these restricted-range species in this publication. However, such information has been presented to relevant government agencies, and is available upon request by fellow scientists.

A new species of the genus Liuixalus (Anura: Rhacophoridae) from southern China.

A new tree frog species of the genus Liuixalus was described from Heishiding Nature Reserve, Guangdong Province, China based on a combination of morphological characters and molecular phylogenetic analyses. Liuixalus feii sp. nov. is distinguished from its congeners by a combination of following characters: small size (SVL 16.2-17.6 mm in adult males and 18.0-18.7 mm in adult females); snout obtusely pointed; tympanum distinct, about half size of eye diameter; nostril closer to eye than to the tip of snout; fingers free of webbing; toe III longer than toe V; toes weakly webbed; tibiotarsal articulation reaching the naris or loreal; dorsal skin smooth and scattered with fine granulars; a subtle longitudinal median ridge present on dorsum; weak skin folds present on dorsal surface of body and thighs; supratympanic fold distinct and curved; ventral surface dull white with more or less irregular dark spots; iris bicolored. The new species appears to be forest-dependent and to date has only been found on the forest floor in primary forests at elevations between 350-800 m. Based on our molecular analyses, we consider Liuixalus catbaensis as a junior synonym of L. calcarius. Thus, with the description of the new species, the genus Liuixalus hitherto contains five recognized species, four of which are endemic to China.

A new species of the genus Gekko Laurenti (Squamata: Sauria: Gekkonidae) from Guangxi, China.

A new species of the genus Gekko is described on the basis of six specimens from Wuming county of Guangxi, southern China. Gekko kwangsiensis sp. nov. is distinguished from other congeners by a combination of the following characters: body relatively small (SVL 64.2-69.7 mm in adults), slender; nares in contact with rostral; internasal absent or single; postmentals two (rarely three), enlarged; interorbital scales between anterior corners of the eyes 29-31; dorsal tubercle rows 9-11; ventral scales between mental and cloacal slit 185-208; midbody scale rows 143-156; ventral scale rows 41-45; subdigital lamellae on first toe 11-13, on fourth toe 13-18; finger and toe webbing weakly developed; tubercles absent on upper surface of fore limbs and hind limbs; precloacal pores nine or ten in males, absent in females; postcloacal tubercle single; tubercles present on dorsal surface of tail base; subcaudals enlarged; dorsal surface of body with 9 or 10 thin light bands between nape and sacrum, and dorsal surface of tail with remarkable black and white bands. Data on the natural history of the new species are provided, and the number of species in the genus Gekko recorded from China is now 17.

A new species of the genus Opisthotropis Gunther, 1872 (Squamata: Colubridae: Natricinae) from Guangdong Province, China.

A new natricid snake of the genus Opisthotropis Gunther, 1872, Opisthotropis laui sp. nov., is described from Mt. Gudou, Jiangmen City, Guangdong Province, China. The new species can be distinguished from other congeners by the combination of the following characters: dorsal scales weakly keeled throughout, in 25:23:23 rows; 10 supralabials; 11 infralabials; two internasals, longer than wide, not touching the loreal; one loreal, not touching the eye; one preocular; two postoculars; one anterior temporal scale; 152 ventrals; 53 subcaudals; body and tail dark olive above, with light yellow crossbars.

A new species of the Gekko japonicus group (Squamata: Sauria: Gekkonidae) from the border region between China and Vietnam.

We describe a new species of the genus Gekko on the basis of 25 specimens from southern China and northern Vietnam. Gekko adleri sp. nov. is distinguished from the remaining congeners by a combination of the following characters: size moderate (SVL < 80 mm); nares in contact with rostral; internasal single, smaller than supranasal; postmentals enlarged; interorbital scales between anterior corners of the eyes 27-36; dorsal tubercle rows 7-11; ventral scales between mental and cloacal slit 168-190; midbody scale rows 123-144; ventral scale rows 35-44; subdigital lamellae on first toe 11-14, on fourth toe 11-15; finger and toe webbing present at base; tubercles absent on upper surface of fore limbs; tubercles on tibia 0-8; precloacal pores 17-21 in males; postcloacal tubercle single; tubercles present on dorsal surface of tail base; subcaudals enlarged; dorsal surface of body with four or five narrow light bands between shoulder and sacrum.