Research Article
Print
Research Article
A new species of the genus Leptobrachella Smith, 1935 (Anura, Megophryidae) from northwestern Guangxi, China
expand article infoZhong Huang‡, Zhao-Chi Zeng§, Jian Wang§, Hao-Tian Wang|, Zhen-Hai Deng, Zhi-Wei Wang#, Xiao-Wen Liao‡, Yun-Ming Mo‡
‡ Natural History Museum of Guangxi, Nanning, China
§ Guangdong Polytechnic of Environmental Protection Engineering, Foshan, China
| Hainan Normal University, Haikou, China
¶ Administration of Guangxi Yachang Orchid National Nature Reserve, Leye, China
# Guangxi Forest Inventory & Planning Institute, Nanning, China

Corresponding author: Yun-Ming Mo ( moyunming@163.com )

Academic editor: Zeeshan Mirza
© 2025 Zhong Huang, Zhao-Chi Zeng, Jian Wang, Hao-Tian Wang, Zhen-Hai Deng, Zhi-Wei Wang, Xiao-Wen Liao, Yun-Ming Mo.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Huang Z, Zeng Z-C, Wang J, Wang H-T, Deng Z-H, Wang Z-W, Liao X-W, Mo Y-M (2025) A new species of the genus Leptobrachella Smith, 1935 (Anura, Megophryidae) from northwestern Guangxi, China. Herpetozoa 38: 321-332. https://doi.org/10.3897/herpetozoa.38.e165088
ZooBank: urn:lsid:zoobank.org:pub:AA5C609D-0234-44DE-8918-CABBD90CB8A1
Open Access

Abstract

In this study, we describe a new species of the species-rich genus Leptobrachella, Leptobrachella yachangensis sp. nov., from northwestern Guangxi, China, based on a combination of molecular and morphological data. The new species is a sister taxon to L. wumingensis Chen, Peng, Li & Yu, 2023, with discrete, diagnostic differences from all other congeners. We also discussed the phenomenon and causes of the high diversity within this genus as well as the conservation of its congeners.

Key Words

conservation, diversity, morphology, molecular phylogeny, taxonomy

Introduction

The genus Leptobrachella Smith, 1925 represents one of the most species-rich groups of frogs with 110 recognized species (Frost 2025). Congeners of this genus are forest dwellers that are widely distributed in China, Myanmar,northeastern India, the Indochina region, Borneo and NatunaIsland (Frost 2025). The morphological conservativeness hinders our understanding of the true diversity of this group (Wang et al. 2019; Wang et al. 2022).

During recent field surveys in mountainous areas of northwestern Guangxi (Fig. 1), we collected seven adult male and two adult female specimens of a Leptobrachella species. Preliminary morphological examination indicated that they can be distinguished from recognized congeners by a series of discrete features. Subsequent molecular analysis further revealed that these specimens represent a separate evolutionary lineage. Considering both the morphological differences and their genetic allocation and divergences, these specimens are described herein as a new species.

Figure 1. 

Distribution sites of Leptobrachella yachangensis sp. nov., L. damingshanensis and L. wumingensis.

Materials and methods

Sampling

During our field work in northwestern Guangxi, three Leptobrachella individuals were collected from Yachang Orchid National Nature Reserve. All newly collected specimens were euthanized with MS-222, fixed in 10% buffered formalin, later transferred to 70% ethanol for long-term preservation, and deposited at the Natural History Museum of Guangxi (NHMG), Nanning City, Guangxi, China. Muscle tissue samples collected from thigh were preserved in 95% ethanol for molecular analysis.

Phylogenetic analyses

DNA extraction, PCR, and sequencing for the three newly collected samples followed Wang et al. (2020). The 16S ribosomal RNA mitochondrial gene (16S rRNA) fragment of 509 base pairs was sequenced for new samples.In total, 74 samples of 64 Leptobrachella species from China along with two outgroup samples were used in this study, encompassing three newly sequenced individuals and others downloaded from GenBank. Detailed information for all samples is given in Suppl. material 1: table S1.

Sequences were aligned using Clustal X 2.0 (Thompson et al. 1997) with defaultparameters. We ran Jmodeltest v2.1.2 (Darriba et al. 2012; with Akaike and Bayesian information criteria) on our alignment and obtained the best-fitting nucleotide substitution model of GTR + I + G (General Time Reversible model, with variable sites modelled according to the Gamma distribution, and a portion of the sites invariant). Phylogenetic analysis was conducted using Bayesian inference (BI) in MrBayes 3.2.4 (Ronquist et al. 2012) and maximum likelihood (ML) in maximum likelihood (ML) in Randomized Axelerated Maximum Likelihood (RAxML, Stamatakis 2006) with RAxML GUI 1.3 (Silvestro and Michalak 2012). For the ML analysis, an optimal tree was obtained and branch supports were evaluated with 1000 rapid bootstrapping replicates. For the BI analysis, two independent runs with four Markov Chain Monte Carlo simulations were performed for ten million iterations and sampled every 1000 iterations. The first 25% of the samples were discarded as burn-in, leaving 7500 samples in the final summary. Convergence of the Markov Chain Monte Carlo simulations was assessed by PSRF < 0.01 and ESS (effective sample size) value > 200 using Tracer 1.4 (http://tree.bio.ed.ac.uk/software/tracer/). Nodes having ML bootstrap values (BS) ≥ 70 and BI posterior probabilities (BPP) ≥ 0.90 were considered well supported. Genetic distances among all Leptobrachella samples were calculated in MEGA 11 using the uncorrected p-distance model with complete deletion of for gaps ormissing data.

Morphological examination

Following Fei et al. (2009) and Rowley et al. (2013), measurements were taken with digital calipers from preserved specimens (Neiko 01407A Stainless Steel 6-Inch Digital Calipers) to the nearest 0.1 mm. Measurements include: snout-vent length (SVL) from the tip of the snout to posterior margin of the vent; head length (HDL) from the tip of the snout to the articulation of the jaw; head width (HDW) at the commissure of the jaws; snout length (SNT) from the tip of the snout to the anterior corner of the eye; eye diameter (EYE) from the anterior corner of the eye to posterior corner of the eye; internasal distance from inner margins of nostrils (IND); interorbital distance (IOD); horizontal diameter of tympanum (TMP); tympanum-eye distance (TEY) from the anterior edge of tympanum to posterior corner of the eye; tibial length (TIB) from the outer surface of the flexed knee to the heel; manus length (ML) from the tip of the third finger to the proximal edge of inner palmar tubercle; pes length (PE) from the tip of the fourth toe to the proximal edge of the inner metatarsal tubercle; length of lower arm and hand (LAHL) from the tip of the third finger to elbow; and hindlimb length (HLL) from the tip of the fourth toe to vent.

Sex was determined by the presence of vocal sac (and vocal sac openings) in males (vs. absence in females). Comparative morphological data for other congeneric species of Leptobrachella were obtained from the literature (Table 1).

Table 1.
Download as
CSV
XLSX

Data source of the currently known species of the genus Leptobrachella.

ID Leptobrachella species Literature obtained
1 L. aerea (Rowley, Stuart, Richards, Phimmachak & Sivongxay, 2010c) Rowley et al. 2010c
2 L. albomarginataWu, Yu, Kilunda, Murphy & Che, 2025 Wu et al. 2025
3 L. aspera Wang, Lyu, Qi & Wang, 2020 Wang et al. 2020
4 L. alpina (Fei, Ye & Li, 1990) Fei et al. 1990; Fei et al. 2009, 2016
5 L. applebyi (Rowley & Cao, 2009) Rowley and Cao 2009
6 L. arayai (Matsui, 1997) Matsui 1997
7 L. ardens (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) Rowley et al. 2016
8 L. aurantirosea Ninh, Nguyen, Le, Nguyen, Quoc, Orlov, Bezman-Moseyko, Le, Nguyen & Ziegler, 2024 Ninh et al. 2024
9 L. batxatensis Hoang, Pham, Phan, Do, Wang, Jiang & Nguyen, 2025 Hoang et al. (2025)
10 L. baluensis Smith, 1931 Dring 1983; Eto et al. 2016
11 L. bashaensis Lyu, Dai, Wei, He, Yuan, Shi, Zhou, Ran, Kuang, Guo, Wei & Yuan, 2020 Lyu et al. 2020a
12 L. bijie Wang, Li, Li, Chen & Wang, 2019 Wang et al. 2019, 2022; Wang et al. 2021
13 L. bidoupensis (Rowley, Le, Tran & Hoang, 2011) Rowley et al. 2011
14 L. bondangensis Eto, Matsui, Hamidy, Munir & Iskandar, 2018 Eto et al. 2018
15 L. botsfordi (Rowley, Dau & Nguyen, 2013) Rowley et al. 2013
16 L. bourreti (Dubois, 1983) Ohler et al. 2011
17 L. brevicrus Dring, 1983 Dring 1983; Eto et al. 2015
18 L. crocea (Rowley, Hoang, Le, Dau & Cao, 2010) Rowley et al. 2010a
19 L. chishuiensis Li, Liu, Wei & Wang, 2020 Li et al. 2020
20 L. damingshanensis Chen, Yu, Cheng, Meng, Wei, Zhou & Lu, 2021 Chen et al. 2021b
21 L. dayaoshanensis Chen, Yu, Meng & Qin, 2024 Chen et al. 2024
22 L. dong Liu, Shi, Li, Zhang, Xiang, Wei & Wang, 2023 Liu et al. 2023
23 L. dorsospina Wang, Lyu, Qi & Wang, 2020 Wang et al. 2020
24 L. dringi (Dubois, 1987) Inger et al. 1995; Matsui and Dehling2012
25 L. dushanensis Li, Li, Cheng, Liu, Wei, Wang, 2024 Li et al. 2024
26 L. duyenae Hoang, Pham, Phan, Do, Wang, Jiang & Nguyen, 2025 Hoang et al. 2025
27 L. eos (Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011) Ohler et al. 2011
28 L. feii Chen, Yuan & Che, 2020 Chen et al. 2020
29 L. firthi (Rowley, Hoang, Dau, Le & Cao, 2012) Rowley et al. 2012
30 L. fritinniens (Dehling & Matsui, 2013) Dehling and Matsui 2013
31 L. fuliginosa (Matsui, 2006) Matsui 2006
32 L. flaviglandulosa Chen, Wang & Che, 2020 Chen et al. 2020
33 L. fusca Eto, Matsui, Hamidy, Munir & Iskandar, 2018 Eto et al. 2018
34 L. gracilis (Günther, 1872) Günther 1872; Dehling 2012b
35 L. graminicola Nguyen, Tapley, Nguyen, Luong & Rowley, 2021 Nguyen et al. 2021
36 L. guinanensis Chen, Li, Peng & Liu, 2024 Chen et al. 2024
37 L. hamidi (Matsui, 1997) Matsui 1997
38 L. heteropus (Boulenger, 1900) Boulenger 1900
39 L. huynhi Hoang, Luong, Nguyen, Nguyen, Ninh, Le, Ziegler & Pham, 2024 Hoang et al. 2024
40 L. isos (Rowley, Stuart, Neang, Hoang, Dau, Nguyen & Emmett, 2015) Rowley et al. 2015a
41 L. itiokai Eto, Matsui & Nishikawa, 2016 Eto et al. 2016
42 L. juliandringi Eto, Matsui & Nishikawa, 2015 Eto et al. 2015
43 L. jinshaensis Cheng, Shi, Li, Liu, Li & Wang, 2021 Cheng et al. 2021
44 L. jinyunensis Shi, Shen, Wang, Jiang & Wang, 2023 Shi et al. 2023
45 L. kajangensis (Grismer, Grismer & Youmans, 2004) Grismer et al. 2004
46 L. kalonensis (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) Rowley et al. 2016
47 L. kecil (Matsui, Belabut, Ahmad & Yong, 2009) Matsui et al. 2009
48 L. khasiorum (Das, Tron, Rangad & Hooroo, 2010) Das et al. 2010
49 L. korifi Matsui, Panha & Eto, 2023 Matsui et al. 2023
50 L. lateralis (Anderson, 1871) Anderson 1871; Humtsoe et al. 2008
51 L. laui (Sung, Yang & Wang, 2014) Sung et al. 2014
52 L. liui (Fei & Ye, 1990) Fei et al. 1990; Fei et al. 2009; Sung et al. 2014; Wang et al. 2022
53 L. macrops (Duong, Do, Ngo, Nguyen & Poyarkov, 2018) Duong et al. 2018
54 L. maculosa (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) Rowley et al. 2016
55 L. mangshanensis (Hou, Zhang, Hu, Li, Shi, Chen, Mo & Wang, 2018) Hou et al. 2018; Wang et al. 2022
56 L. maoershanensis (Yuan, Sun, Chen, Rowley & Che, 2017) Yuan et al. 2017
57 L. marmorata (Matsui, Zainudin & Nishikawa, 2014b) Matsui et al. 2014b
58 L. maura (Inger, Lakim, Biun & Yambun, 1997) Inger et al. 1997
59 L. melanoleuca (Matsui, 2006) Matsui 2006
60 L. melica (Rowley, Stuart, Neang & Emmett, 2010) Rowley et al. 2010b
61 L. minima (Taylor, 1962) Taylor 1962; Ohler et al. 2011
62 L. mjobergi Smith, 1925 Eto et al. 2015
63 L. murphyi Chen, Suwannapoom, Wu, Poyarkov, Xu, Pawangkhanant & Che, 2021 Chen et al. 2021a
64 L. nahangensis (Lathrop, Murphy, Orlov & Ho, 1998) Lathrop et al. 1998
65 L. natunae (Günther, 1895) Günther 1895
66 L. namdongensis Hoang, Nguyen, Luu, Nguyen & Jiang, 2019 Hoang et al. 2019
67 L. neangi Stuart & Rowley, 2020 Stuart and Rowley 2020
68 L. niveimontis Chen, Poyarkov, Yuan & Che, 2020 Chen et al. 2020
69 L. nokrekensis (Mathew & Sen, 2010) Mathew and Sen 2010
70 L. nyx (Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011) Ohler et al. 2011
71 L. oshanensis (Liu, 1950) Liu, 1950; Shi et al. 2021
72 L. pallida (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) Rowley et al. 2016
73 L. palmata Inger & Stuebing, 1992 Inger and Stuebing 1992
74 L. parva Dring, 1983 Dring 1983
75 L. pelodytoides (Boulenger, 1893) Boulenger 1893; Ohler et al. 2011
76 L. petrops (Rowley, Dau, Hoang, Le, Cutajar & Nguyen, 2017) Rowley et al. 2017a
77 L. phiadenensis Luong, Hoang, Pham, Ziegler & Nguyen, 2023 Luong et al. 2023
78 L. phiaoacensis Luong, Hoang, Pham, Ziegler & Nguyen, 2023 Luong et al. 2023
79 L. pingbianensis (Rao, Hui, Zhu & Ma, 2022 “2020”) Rao et al. 2022 “2020”
80 L. picta (Malkmus, 1992) Malkmus 1992
81 L. platycephala (Dehling, 2012) Dehling 2012a
82 L. pluvialis (Ohler, Marquis, Swan & Grosjean, 2000) Ohler et al. 2000, 2011
83 L. puhoatensis (Rowley, Dau & Cao, 2017) Rowley et al. 2017b
84 L. purpurus (Yang, Zeng & Wang, 2018) Yang et al. 2018
85 L. purpuraventra Wang, Li, Li, Chen & Wang, 2019 Wang et al. 2019
86 L. pyrrhops (Poyarkov, Rowley, Gogoleva, Vassilieva, Galoyan & Orlov, 2015) Poyarkov et al. 2015
87 L. rowleyae (Nguyen, Poyarkov, Le, Vo, Ninh, Duong, Murphy & Sang, 2018) Nguyen et al. 2018
88 L. sabahmontana (Matsui, Nishikawa & Yambun, 2014) Matsui et al. 2014a
89 L. serasanae Dring, 1983 Dring 1983
90 L. shangsiensis Chen, Liao, Zhou & Mo, 2019 Chen et al. 2019
91 L. shimentaina Wang, Lyu & Wang, 2022 Wang et al. 2022
92 L. sinorensis Matsui, Panha & Eto, 2023 Matsui et al. 2023
93 L. sola (Matsui, 2006) Matsui 2006
94 L. suiyangensis Luo, Xiao, Gao & Zhou, 2020 Luo et al. 2020
95 L. sungi (Lathrop, Murphy, Orlov & Ho, 1998) Lathrop et al. 1998
96 L. shiwandashanensis Chen, Peng, Pan, Liao, Liu & Huang, 2021 Chen et al. 2021c
97 L. tadungensis (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) Rowley et al. 2016
98 L. tamdil (Sengupta, Sailo, Lalremsanga, Das & Das, 2010) Sengupta et al. 2010
99 L. tengchongensis (Yang, Wang, Chen & Rao, 2016) Yang et al. 2016
100 L. tuberosa (Inger, Orlov & Darevsky, 1999) Inger et al. 1999
101 L. ventripunctata (Fei, Ye & Li, 1990) Fei et al. 1990; Fei et al. 2009, 2016
102 L. verrucosa Wang, Zeng, Lin & Li, 2022 Lin et al. 2022
103 L. wuhuangmontis Wang, Yang & Wang, 2018 Wang et al. 2018
104 L. wulingensis Qian, Xia, Cao, Xiao & Yang, 2020 Qian et al. 2020
105 L. wumingensis Chen, Peng, Li & Yu, 2023 Chen et al. 2023
106 L. xishuiensis Luo, Deng, Zhou, 2025 Luo et al. 2025
107 L. yingjiangensis (Yang, Zeng & Wang, 2018) Yang et al. 2018
108 L. yongshunensis Huang, Wu, Jiang & Zhang, 2025 Huang et al. 2025
109 L. yunkaiensis Wang, Li, Lyu & Wang, 2018 Wang et al. 2018
110 L. yeae Shi, Hou, Song, Jiang & Wang, 2021 Shi et al. 2021
111 L. yunyangensis Luo, Deng & Zhou, 2022 Luo et al. 2022
112 L. zhangyapingi (Jiang, Yan, Suwannapoom, Chomdej & Che, 2013) Jiang et al. 2013

Results

The ML and Bayesian analysis both resulted identical topologies and strongly support that all Leptobrachella samples formed a monophyletic clade (Fig. 2). In the phylogenetic topology, the Leptobrachella samples from Yachang Orchid National Nature Reserve form a monophyletic lineage with strong support (BS = 100; BPP = 1.00). Though with low support, both ML and Bayesian analysis result that the new lineage is a sister taxon to L. wumingensis. Besides, the new lineage from Yachang Orchid National Nature Reserve obviously differs from its closely-related congeners by mean p-distance ≥ 6.5% in the 16S RNA ribosomal mitochondrial gene, which significantly exceeds the level of genetic divergence observed in recognized species of Leptobrachella.

Figure 2. 

Maximum Likelihood tree results from partial DNA sequences of the mitochondrial 16S r RNA gene. Numbers before slashes indicate bootstrap support (BS) and numbers after slashes are Bayesian posterior probabilities (BPP). The ‘-’ represent BS ≤ 70 or BPP ≤ 0.90. Voucher before species name at the branch terminal corresponds to the voucher ID in Suppl. material 1: table S1.

Our detailed morphological examination has also revealed discrete, diagnostic (non-overlapping ranges in traditional characters) differences among the specimens from this independent lineage and all other congeners. Therefore, both the phylogenetic result and morphological comparison support the hypothesis that the lineage from Yachang Orchid National Nature Reserve represent an undescribed new species, and herein we describe this new species.

Taxonomic account

Leptobrachella yachangensis sp. nov.

https://zoobank.org/87019B8E-76B9-4738-9BCD-1876D40C40F4
Fig. 3 Yachang Leaf Litter Toad (in English) / Ya Chang Zhang Tu Chan (雅长掌突蟾 in Chinese)

Type materials.

Holotype ♂.NHMG250402, collected by Yun-Ming Mo and Zhong Huang on 15 April 2025 from Guangxi Yachang Orchid National Nature Reserve (24°44'57.24" N, 106°19'38.94" E; ca. 1400 m a.s.l.), Leye County, Baise City, Guangxi, China.

Figure 3. 

Morphological features of the holotype of Leptobrachella yachangensissp. nov., NHMG250402 in life. A. Dorsal view; B. Dorsolateral view; C. Lateral view; D. Ventral view; E. Ventral view of hand; F. Ventral view of foot.

Paratypes. 6 ♂: NHMG250401,NHMG250409-250414, the same collection data as the holotype; NHMG250401, collected by Yun-Ming Mo and Zhong Huang on 15 April 2025 from Guangxi Yachang Orchid National Nature Reserve (24°45'6.85" N, 106°19'22.38" E; ca. 1250 m a.s.l.); NHMG250409-250414, collected by Yun-Ming Mo and Zhong Huang on 16 April 2025 from Guangxi Yachang Orchid National Nature Reserve (24°44'52.82"N, 106°20'2.06"E; ca. 1610 m a.s.l.).

Etymology.

The specific epithet, yachangensis, is in reference to the type locality, Yachang Orchids National Nature Reserve.

Diagnosis.

(1) Small body size [SVL 28.1–31.2 mm in seven adult males], (2) iris bicolored, upper half coppery orange and lower half grayish brown, (3) tympanum distinct, (4) supratympanic line black, (5) fingers without lateral fringes, (6) toes with narrow lateral fringes,lacking webbing, (7) longitudinal ridges under toes continuous, (8) heels overlapping when adpressed, tibial-tarsal articulation reaching anterior corner of eye, (9) dorsal surface rough with dense skin ridges and numerous conical tubercles bearing spines, (10) ventral surface smooth, with transparent tiny conical spines on chest and abdomen in breeding males, (11) dorsum grayish brown, with darker brown scattered markings, (12) flanks with several dark spots, (13) surface of throat, chest and abdomen creamy white, with dense irregular dark brown speckles.

Description of holotype.

Adult male. Body size small, SVL 31.2 mm. Head length slightly larger than head width, HDW/HDL 0.90; snout slightly protruding, projecting slightly beyond margin of lower jaw; nostril closer to snout than eye; canthus rostralis gently rounded; loreal region slightly concave; interorbital space flat, internarial distance slightly larger than interorbital distance, IND/IOD 1.07; pineal ocellus absent; pupil vertical; snout slightly longer than eye diameter, SNT/EYE 1.07; tympanum distinct, rounded, diameter smaller than that of eye and larger than tympanum-eye distance, TMP/EYE 0.48, TEY/TMP 0.48; upper margin of tympanum in contact with supratympanic ridge; vomerine teeth absent; a single vocal sac; vocal sac openings slit-like, paired, located posterolaterally on floor of mouth, close to margins of mandible; tongue deeply notched distally; supratympanic ridge distinct, extending from posterior corner of eye to supra-axillary gland.

Tips of fingers rounded, slightly swollen; relative finger lengths I < II < IV < III; nuptial pad absent; subarticular tubercles absent; inner palmar tubercle large, rounded, distinctly separated from outer palmar tubercle; outer palmar tubercle small, rounded; fingers lacking interdigital webbing and lateral fringes. Tips of toes rounded, slightly swollen; relative toe length I < II < V< III < IV; longitudinal ridges under toes continuous; inner metatarsal tubercle large, oval; outer metatarsal tubercle absent; interdigital webbing between toes absent; narrow lateral fringes present on all toes. Tibia 47% of snout-vent length; tibiotarsal articulation reaches to anterior corner of eye; heels overlapping when thighs are appressed at right angles with respect to body.

Dorsal surface rough with dense skin ridges and numerous conical tubercles bearingspines; ventral skin smooth with transparent tiny conical spines on chest and abdomen; pectoral and femoral glands oval, both larger in diameter than tip of fingers; femoral glands larger in diameter than tip of toes; femoral gland situated on posteroventral surface of thigh, closer to knee than to vent; supra-axillary gland raised. Ventrolateral gland distinctly visible, forming a longitudinal uncontinuous series.

Coloration of holotype in life.

Dorsal background color grayish brown, with darker brown markings. A dark brown inverted triangular marking in interorbital region, connecting to dark brown W-shaped marking on occipital region. Tympanum grayish brown. Supratympanic line black. Transverse dark brown bars on dorsal surfaces of distal limbs and digits; distinct dark brown spots on flanks; surfaces of digits, elbows, upper arms and tibiotarsal articulations coppery orange, with dark markings and spots.

Surface of throat, chest and abdomen creamy white, with dense irregular dark brown speckles. Underside of limbsgrayish brown with numerous irregular white speckles. Supra-axillary gland coppery orange, pectoral and femoral glands grayish white, ventrolateral glands grayish white. Iris bicolored, upper half coppery orange and lower half grayish brown.

Coloration of holotype in preservative.

Dorsal background color dark brown scattered with irregular gray pigmentations. All markings, bars and spots become indistinct. Tympanum dark brown.

Ventral surface of throat, chest and abdomen grayish white; irregular dark brown speckles more distinct. Underside of limbs dark brown, irregular speckles grayish white. Supra-axillary, pectoral, and ventrolateral glands grayish white.

Variation.

Measurements and body proportions are listed in Table 2. All the specimens showed relatively small differences in coloration and color patterns.

Table 2.
Download as
CSV
XLSX

Measurements (minimum–maximum (mean ± SD); in mm), and body proportions ofLeptobrachella yachangensis sp. nov.

Vocher NHMG 250401 NHMG 250402 (Holotype) NHMG 250409 NHMG 250411 NHMG 250412 NHMG 250413 NHMG 250414 Range
Sex Male Male Male Male Male Male Male Males (n = 7)
SVL 28.3 31.2 30.0 28.7 28.9 28.1 29.8 28.1–31.2 (29.3 ± 1.1)
HDL 11.1 11.7 11.4 11.0 10.8 11.5 11.2 10.8–11.7 (11.2 ± 0.3)
HDW 10.1 10.5 10.1 10.0 10.4 10.3 11.6 10.0–11.6 (10.4 ± 0.5)
SNT 4.4 4.7 4.5 4.4 4.4 4.6 4.5 4.4–4.7 (4.5 ± 0.1)
IND 2.8 3.1 3.0 2.9 3.1 2.9 3.1 2.8–3.1 (3.0 ± 0.1)
IOD 2.7 2.9 2.8 2.7 2.8 2.6 2.8 2.6–2.9 (2.8 ± 0.1)
EYE 4.2 4.4 4.1 4.1 4.2 4.3 4.5 4.1–4.5 (4.3 ± 0.2)
TMP 1.9 2.1 2.0 2.1 2.0 2.1 2.1 1.9–2.1 (2.0 ± 0.1)
TEY 1.0 0.7 0..8 0.8 0.9 0.8 0.9 0.7–1.0 (0.9 ± 0.1)
ML 8.6 8.2 8.3 8.5 8.2 8.3 8.4 8.2–8.6 (8.4 ± 0.2)
LAHL 16.2 15.4 15.7 16.0 15.5 15.8 15.9 15.4–16.2 (15.8 ± 0.3)
PL 14.7 13.7 14.3 14.6 14.2 13.8 14.4 13.7–14.7 (14.2 ± 0.4)
TIB 14.3 14.7 14.1 14.9 14.5 14.2 14.8 14.1–14.9 (14.5 ± 0.3)
HLL 42.7 46.8 44.6 46.3 45.7 44.4 45.9 42.7–46.8 (45.2 ± 1.4)

Distribution, ecology and conservation.

Leptobrachella yachangensis sp. nov. is currently only known from its type locality (Fig. 1; ca. 1240–1620 m a.s.l.). The new species inhabits clear-water rocky streams in primary forests surrounded by broad-leaved forest in granite landforms of karst region. Males were observed calling while perched in the rock crevices or on the leaves of dwarf shrubs near flowing seeps. The breeding season lasts from February to June according to our long-term observation. During the breeding season, a large number of vocalizing individuals can be observed in the streams.Leptobrachella yachangensis sp. nov.is under multiple potential human-induced pressures: potential wind power projects and eco-tourism development outside the protected area, artificial forest (such as Chinese fir forest) planting in the surrounding areas, and other activities may cause water pollution and habitat fragmentation. However, given that the current information on the distribution and threats of this species is still not fully understood, we recommend the new species be listed as Data Deficient (DD) according to the International Union for Conservation of Nature (IUCN) Red List Categories and Criteria.

Comparisons.

In the phylogenetic tree (Fig. 2),Leptobrachella yachangensis sp. nov. is most closely related to L. wumingensis (mean p-distance 6.5%) and L. damingshanensis (mean p-distance 7.2%), therefore, these two species are selected for detailed comparison. The new species differs from L. wumingensis by its presence of narrow fringes along lateral surfaces of toes (vs. absence of lateral fringes on toes), and differs from L. damingshanensis by its absence of webbing on toes (vs. presence of webbing on toes). Moreover, the new species can be also distinguished from the later two species by its presence of numerous conical tubercles bearing spines on dorsal surface (vs. absence of spines), the presence of transparent tiny conical spines on chest and abdomen in breeding males (vs. absence of spines), presence of dense irregular dark brown speckles on throat, chest and abdomen with (vs. absence of such speckles).

Compared with the 26 known congeners of the genus Leptobrachella occurring south of the Isthmus of Kra, by the presence of supra-axillary and ventrolateral glands, L. yachangensis sp. nov. can easily be distinguished from L. arayai, L. dringi, L. fritinniens, L. gracilis, L. hamidi, L. heteropus, L. kajangensis, L. kecil, L. marmorata, L. melanoleuca, L. maura, L. picta, L. platycephala, L. sabahmontana, and L. sola, all of which lack supra-axillary and ventrolateral glands; and by its distinctly larger male body size, SVL 25.7–28.2 mm, L. yachangensis sp. nov. differs from the smaller L. baluensis (SVL 14.9–15.9 mm), L. brevicrus (SVL 17.1–17.8 mm), L. bondangensis (SVL 17.8 mm), L. fusca (SVL 16.3 mm), L. itiokai (SVL 15.2–16.7 mm), L. juliandringi (SVL 17.0–17.2 mm), L. mjobergi (SVL 15.7–19.0 mm), L. natunae (SVL 17.6 mm), L. parva (SVL 15.0–16.9 mm), and L. palmata (SVL 14.4–16.8 mm); and even distinctly larger than female L. serasanae (SVL 16.9 mm).

In having black spots on flanks, Leptobrachella yachangensis sp. nov. further differs from L. aerea, L. botsfordi, L. crocea, L. eos, L. firthi, L. isos, L. pallida, L. petrops, and L. tuberosa, all of which lack black spots on the flanks.

For the remaining 75 members of the genus Leptobrachella, males of the new species (SVL 28.1–31.2 mm) differ from males of the larger L. nahangensis (40.8 mm), L. sungi (48.3–52.7 mm), and L. zhangyapingi (48.5–52.5 mm); and males of the smaller L. applebyi (19.6–22.3 mm), L. aspera (22.4 mm), L. feii (21.5–22.8 mm), L. melica (19.5–22.7 mm), L. niveimontis (22.5–23.6 mm), L. pluvialis (21.3–22.3 mm), and even distinctly larger than female L. korifi (22.7 mm).

Through its toes without webbing, Leptobrachella yachangensis sp. nov. can be distinguished from L. albomarginata, L. alpina, L. bashaensis, L. bijie, L. bidoupensis, L. bourreti, L. chishuiensis, L. dayaoshanensis, L. dong, L. dorsospina, L. dushanensis, L. guinanensis, L. fuliginosa, L. flaviglandulosa, L. graminicola, L. jinyunensis, L. khasiorum, L. lateralis, L. laui, L. liui, L. macrops, L. mangshanensis, L. maoershanensis, L. minima, L. murphyi, L. namdongensis, L. neangi, L. nokrekensis, L. nyx, L. pelodytoides, L. puhoatensis, L. purpurus, L. purpuraventra, L. pyrrhops, L. phiadenensis, L. phiaoacensis, L. pingbianensis, L. rowleyae, L. shangsiensis, L. shimentaina, L. sinorensis, L. suiyangensis, L. tamdil, L. tengchongensis, L. ventripunctata, L. verrucosa, L. wulingensis, L. xishuiensis, L. yongshunensis, L. yingjiangensis, L. yunkaiensis, L. yunyangensis, and L. yeae, all of which bear webbing on toes. In having toes with narrow lateral fringes, the new species can be distinguished from L. ardens, L. kalonensis, L. maculosa, L. oshanensis, L. shiwandashanensis, and L. tadungensis (vs. lateral fringes absent). In having transparent tiny conical spines on chest and abdomen in breeding males, the new species can be distinguished from L. aurantirosea, L. batxatensis, L. huynhi, and L. jinshaensis (vs. ventral skin smooth).

Discussion

The rapid discovery of numerous cryptic species within the genus Leptobrachella across southern China in the past five years (37 new species described since 2020, 10 of which are from Guangxi; Chen et al. 2020; Wang et al. 2022; Liu et al. 2023; Li et al. 2024) strongly indicates that the true diversity of this genus remains severely underestimated. Our discovery of the sympatric occurrence of Leptobrachella yachangensis and L. ventripunctata in rocky stream habitats within the karst mountains of Leye County, Guangxi (bordering Guizhou), further corroborates the findings of Rowley et al. (2015): highly divergent lineages within Leptobrachella frequently exhibit pronounced micro-endemism and sympatry. This complex distribution pattern significantly complicates species delimitation and classification within the genus. Furthermore, the widespread morphological conservatism among Leptobrachella species (Wang et al. 2019; Chen et al. 2020), resulting in highly similar appearances of many congeners in the field, not only hinders research into their diversification mechanisms but also poses challenges for developing targeted conservation strategies.

Molecular phylogenetic analysis (Fig. 2) revealed that L. yachangensis is sister to L. wumingensis, not to the sympatrically occurring L. ventripunctata. This phenomenon of "closely related species being allopatric, while distantly related species co-occurring sympatrically" is not unique within Leptobrachella. For instance: in Guangdong Shimentai National Nature Reserve, L. shimentaina co-occurs with the phylogenetically distant L. mangshanensis (Wang et al. 2022); in Yingjiang County of Yunnan, L. purpurus is found sympatrically with the phylogenetically distant L. yingjiangensis and L. ventripunctata (Yang et al. 2018). These cases collectively demonstrate a prevalent pattern of "decoupling of phylogeny and sympatry" within Leptobrachella, suggesting that niche differentiation (e.g., microhabitat preference, breeding season isolation) or historical dispersal events may play a more critical role than shared ancestry in driving sympatric distribution patterns (Rowley et al. 2015). Geographically, the only currently known locality of L. yachangensis (Yachang Orchid NNR, Leye, Guangxi) is approximately 250 km in a straight line from the type locality of its sister species, L. wumingensis (Mt Damingshan, Wuming, Guangxi), separated by the Hongshui River valley and continuous non-karst mountains. This moderate-scale geographic isolation, combined with significant genetic divergence (p-distance 4.1–4.9%), supports the hypothesis of karst landscapes acting as "evolutionary islands" (Ficetola et al. 2021), where isolated habitat patches drive the independent evolution of genetic lineages.

Leptobrachella yachangensis is currently known only from primary montane forests at elevations of 1240–1620 m within the Yachang Orchid National Nature Reserve, Leye County, Guangxi. Its survival is under multiple potential human-induced pressures: potential wind power projects and eco-tourism development outside the protected area, artificial forest (such as Chinese fir forest) planting in the surrounding areas, and other activities may cause water pollution and habitat fragmentation. Climate change may also affect the micro-environment of the alpine streams it depends on. Therefore, we recommend the immediate implementation of the following conservation measures: 1) Expand survey efforts: Systematically survey adjacent karst regions (e.g., Maolan NNR in Guizhou, Mulun NNR in Guangxi) to assess the species’ actual distribution range and population connectivity; 2) Strengthen habitat protection: Strictly limit anthropogenic disturbances (e.g., trail construction, tourist access) within core stream habitats in the reserve and maintain the integrity of the primary forest vegetation; 3) Establish dynamic monitoring: Implement a population and habitat monitoring network to evaluate the long-term impacts of tourism and quarrying activities. Given its extremely limited distribution and evident threats, and although current data is insufficient for a precise assessment, we strongly recommend its inclusion in the China Biodiversity Red List and prioritization for an IUCN Red List assessment in the near future (adopting the precautionary principle from DD towards potentially VU).

Acknowledgements

We thank Ben-Zhe Huang for his help in the field work. This work was supported by the Project of Wildlife Theme 2-Amphibian Reptiles of the Southwest Karst National Park (Guangxi) Comprehensive Scientific Investigation Project (LKWT-2023-095).

References

  • Anderson J (1871) A list of the reptilian accession to the Indian Museum, Calcutta from 1865 to 1870, with a description of some new species. Journal of the Asiatic Society of Bengal 40: 12–39.
  • Boulenger GA (1893) Concluding report on the reptiles and batrachians obtained in Burma by Signor L. Fea dealing with the collection made in Pegu and the Karin Hills in 1887–88. Annali del Museo Civico di Storia Naturale di Genova 13: 304–347.
  • Chen JM, Poyarkov Jr NJ, Suwannapoom C, Lathrop A, Wu YH, Zhou WW, Yuan ZY, Jin JQ, Chen HM, Liu HQ, Nguyen TQ, Nguyen SN, Duong TV, Eto K, Nishikawa K, Matsui M, Orlov NL, Stuart BL, Brown RM, Rowley JJL, Murphy RW, Wang YY, Che J (2018) Large-scale phylogenetic analyses provide insights into unrecognized diversity and historical biogeography of Asian leaf-litter frogs, genus Leptolalax (Anura: Megophryidae). Molecular Phylogenetics and Evolution 124: 162–171. https://doi.org/10.1016/j.ympev.2018.02.020
  • Chen JM, Xu K, Poyarkov NA, Wang K, Yuan ZY, Hou M, Suwannapoom C, Wang J, Che J (2020) How little is known about “the little brown frogs”: description of three new species of the genus Leptobrachella (Anura: Megophryidae) from Yunnan Province, China. Zoological Research 41: 1–22. https://doi.org/10.24272/j.issn.2095-8137.2020.036
  • Chen JM, Suwannapoom C, Wu YH, Poyarkov NA, Xu K, Pawangkhanant P, Che J (2021a) Integrative taxonomy reveals a new species of Leptobrachella (Anura: Megophryidae) from the mountains of northern Thailand. Zootaxa 5052: 191–214. https://doi.org/10.11646/zootaxa.5072.2.8
  • Chen WC, Li P, Peng WX, Liu YJ, Huang Y (2024) The fourth species of Leptobrachella (Anura, Megophryidae) found at Shiwandashan National Nature Reserve, Guangxi, China. ZooKeys 1192: 257–279. https://doi.org/10.3897/zookeys.1192.98352
  • Chen WC, Peng WX, Pan WH, Liao NY, Liu YJ, Huang Y (2021c) A new species of Leptobrachella Smith 1925 (Anura: Megophryidae) from Southern Guangxi, China. Zootaxa 5020(3): 581–596. https://doi.org/10.11646/zootaxa.5020.3.8
  • Cheng YL, Shi SC, Li J, Liu J, Li SZ, Wang B (2021) A new species of the Asian leaf litter toad genus Leptobrachella Smith, 1925 (Anura, Megophryidae) from northwest Guizhou Province, China. ZooKeys 1021: 81–107. https://doi.org/10.3897/zookeys.1021.60729
  • Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: More models, new heuristics and parallel computing. Nature Methods 9(8): 772. https://doi.org/10.1038/nmeth.2109
  • Das I, Tron RKL, Rangad D, Hooroo RN (2010) A new species of Leptolalax (Anura: Megophryidae) from the sacred groves of Mawphlang, Meghalaya, north-eastern India. Zootaxa 2339(1): 44–56. https://doi.org/10.11646/zootaxa.2339.1.2
  • Dehling JM (2012a) Eine neue Art der Gattung Leptolalax (Anura: Megophryidae) vom Gunung Benom, Westmalaysia/A new species of the genus Leptolalax (Anura: Megophryidae) from GunungBenom, Peninsular Malaysia. Sauria 34: 9–21.
  • Dehling JM (2012b) Redescription of Leptolalax gracilis (Günther, 1872) from Borneo and taxonomic status of two populations of Leptolalax (Anura: Megophryidae) from Peninsular Malaysia. Zootaxa 3328(1): 20–34. https://doi.org/10.11646/zootaxa.3328.1.2
  • Dehling JM, Matsui M (2013) A new species of Leptolalax (Anura: Megophryidae) from Gunung Mulu National Park, Sarawak, East Malaysia (Borneo). Zootaxa 3670(1): 33–44. https://doi.org/10.11646/zootaxa.3670.1.2
  • Dubois A (1983) Note préliminaire sur le genre Leptolalax Dubois, 1980 (Amphibiens, Anoures), avec diagnose d’uneespèce novelle du Vietnam. Alytes 2: 147–153. https://doi.org/10.5962/p.297429
  • Dubois A (1987) “1986” Miscellanea taxinomica batrachologica (I). Alytes. Paris 5: 7–95.
  • Eto K, Matsui M, Nishikawa K (2015) Description of a new species of the genus Leptobrachella (Amphibia, Anura, Megophryidae) from Borneo. Current Herpetology 34(2): 128–139. https://doi.org/10.5358/hsj.34.128
  • Eto K, Matsui M, Nishikawa K (2016) A new highland species of dwarf litter frog genus Leptobrachella (Amphibia, Anura, Megophryidae) from Sarawak. The Raffles Bulletin of Zoology 64: 194–203.
  • Eto K, Matsui M, Hamidy A, Munir M, Iskandar DT (2018) Two new species of the genus Leptobrachella (Amphibia: Anura: Megophryidae) from Kalimantan, Indonesia. Current Herpetology 37(2): 95–105. https://doi.org/10.5358/hsj.37.95
  • Fei L, Ye CY (1992) Comments on the taxonomy of pelobatid toads of genus Leptolalax (Carpophrys) with description of a new species. Acta Zoological Sinica 38(3): 245–253. [In Chinese with English abstract]
  • Fei L, Ye CY, Huang YZ (1990) Key to Chinese Amphibians. Publishing House for Scientific and Technological Literature, Chongqing, 364 pp. [In Chinese]
  • Fei L, Hu SQ, Ye CY, Huang YZ (2009) Fauna Sinica. Amphibia Vol. 2 Anura. Science Press, Beijing, 957 pp. [In Chinese]
  • Fei L, Ye CY, Jiang JP (2012) Colored atlas of Chinese amphibians and their distributions. Sichuan Publishing House of Science & Technology, Chengdu, 619 pp. [In Chinese]
  • Grismer LL, Grismer JL, Youmans TM (2004) A new species of Leptolalax (Anura: Megophryidae) from Pulau Tioman, West Malaysia. Asiatic Herpetological Research 10: 8–11.
  • Günther A (1872) On the reptiles and amphibians of Borneo. Proceedings of the Scientific Meetings of the Zoological Society of London 1872: 586–600.
  • Günther A (1895) The reptiles and batrachians of the Natuna Islands. Novitates Zoologicae 2: 499–502.
  • Hoang CV, Nguyen TT, Luu VQ, Nguyen TQ, Jiang JP (2019) A new species of Leptobrachella Smith 1925 (Anura: Megophryidae) from Thanh Hoa Province, Vietnam. Raffles Bulletin of Zoology. Singapore 67: 536–556. https://doi.org/10.26107/RBZ–2019-0042
  • Huang J, Zhang FP, Jiang WS, Tian YX, Huang XL, Xu YL, Liu J, Li XY, Zhang XY, Wu T (2025) Description of a new leaf litter toad of Leptobrachella (Anura, Megophryidae) from Hunan, China. Zoosystematics and Evolution 101(1): 103–117. https://doi.org/10.3897/zse.101.135586
  • Hou YM, Zhang MF, Hu F, Li SY, Shi SC, Chen J, Mo XY, Wang B (2018) A new species of the genus Leptolalax (Anura, Megophryidae) from Hunan, China. Zootaxa 4444(3): 247–266. https://doi.org/10.11646/zootaxa.4444.3.2
  • Hoang CV, Luong AM, Nguyen TQ, Nguyen TT, Ninh HT, Le LTH, Ziegler T, Pham CT (2024) A new species of Leptobrachella Smith 1925 (Anura, Megophryidae) from Lai Chau Province, Vietnam. Biodiversity Data Journal 12(e136491): 1–18. https://doi.org/10.3897/BDJ.12.e136491
  • Hoang VC, Pham TC, Phan QT, Do TD, Wang B, Jiang JP, Nguyen QT (2025) Hidden biodiversity in the tropicalrain forests: Two new species of Leptobrachella Smith 1925 (Anura: Megophryidae) from Vietnam. Asian Herpetological Research 16(2): 161–169. https://doi.org/10.3724/ahr.2095-0357.2024.0039
  • Inger RF, Stuebing RB (1992) “1991”. A new species of frog of the genus Leptobrachella Smith (Anura: Pelobatidae), with a key to the species from Borneo. Raffles Bulletin of Zoology. Singapore 39: 99–103.
  • Inger RF, Stuebing RB, Tan FL (1995) New species and new records of anurans from Borneo. The Raffles Bulletin of Zoology 43: 115–132.
  • Inger RF, Lakim M, Biun A, Yambun P (1997) A new species of Leptolalax (Anura: Megophryidae) from Borneo. Asiatic Herpetological Research 7: 48–50. https://doi.org/10.5962/bhl.part.18855
  • Jiang K, Yan F, Suwannapoom C, Chomdej S, Che J (2013) A new species of the genus Leptolalax (Anura: Megophryidae) from northern Thailand. Asian Herpetological Research 4(2): 100–108. https://doi.org/10.3724/SP.J.1245.2013.00100
  • Lathrop A, Murphy RW, Orlov N, Ho CT (1998) Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Amphibia-Reptilia 19(3): 253–267. https://doi.org/10.1163/156853898X00160
  • Luong AM, Hoang CV, Pham CT, Ziegler T, Nguyen TQ (2023) Two new species of Leptobrachella Smith 1925 (Amphibia: Megophryidae) from Cao Bang Province, Vietnam. Zootaxa 5369(3): 301–335. https://doi.org/10.11646/zootaxa.5369.3.1
  • Li ZC, Xiao Z, Liu SR (2011) Amphibians and Reptiles of Guangdong. Guangdong Science & Technology Press, LTD, Guangzhou, 266 pp. [in Chinese]
  • Li SZ, Liu J, Wei G, Wang B (2020) A new species of the Asian leaf litter toad genus Leptobrachella (Amphibia, Anura, Megophryidae) from southwest China. ZooKeys 943: 91–118. https://doi.org/10.3897/zookeys.943.51572
  • Li SZ, Li W, Cheng YL, Liu J, Wei G, Wang B (2024) Description of a new Asian Leaf Litter Toad of the genus Leptobrachella Smith, 1925 (Anura, Megophryidae) from southern Guizhou Province, China. Biodiversity Data Journal 12(e113427): 1–35. https://doi.org/10.3897/BDJ.12.e113427
  • Lin SS, Li YH, Lu YH, Su HL, Wu SB, Zhang QQ, Mo MJ, Xiao SJ, Pan Z, Pan HJ, Zeng ZC, Wang J (2022) A new species of the genus Leptobrachella (Anura, Megophryidae) from northwestern Guangdong Province, China. Herpetozoa Wien 35: 165–178. https://doi.org/10.3897/herpetozoa.35.e89981
  • Liu J, Shi SC, Li SZ, Zhang MF, Xiang SJ, Wei G, Wang B (2023) A new Asian leaf litter toad of the genus Leptobrachella (Amphibia, Anura, Megophryidae) from central south China. ZooKeys 1149: 103–134. https://doi.org/10.3897/zookeys.1149.85895
  • Luo T, Wang WF, Peng D, Lei B, Deng HQ, Ji SN, Huang HQ, Zhou J (2022) A new species of the asianleaf litter toad genus Leptobrachella (Amphibia,Anura, Megophryidae) from Chongqing city, southwest China. Asian Herpetological Research 13(2): 75–95. https://doi.org/10.16373/j.cnki.ahr.210052
  • Luo T, Zhao ZF, Wang ZL, Xiao MY, Deng H, Xiao N, Zhou J (2025) Diversification outbreaks and dynamics of Asian leaf-litter frogs, genus Leptobrachella (Anura, Megophryidae), with the description of a new species from Guizhou Province, China. Zoosystematics and Evolution 101(1): 223–243. https://doi.org/10.3897/zse.101.137392
  • Lyu ZT, Qi S, Wang J, Zhang SY, Zhao J, Zeng ZC, Wan H, Yang JH, Mo YM, Wang YY (2023) Generic classification of Asian Horned Toads (Anura: Megophryidae: Megophryinae) and monograph of Chinese species. Zoological Research. Kunming 44: 380–450. https://doi.org/10.24272/j.issn.2095-8137.2022.372
  • Lyu JC, Dai LL, Wei PF, He YH, Yuan ZY, Shi WL, Zhou SL, Ran SY, Kuang ZF, Guo X, Wei G, Yuan G (2020a) A new species of the genus Leptobrachella Smith, 1925 (Anura, Megophryidae) from Guizhou, China. ZooKeys 1008: 139–157. https://doi.org/10.3897/zookeys.1008.56412
  • Malkmus R (1992) Leptolalax pictus sp.n. (Anura: Pelobatidae) vom Mount Kinabalu/Nord-Borneo. Sauria 14: 3–6.
  • Mathew R, Sen N (2010) [2009] Description of a new species of Leptobrachium Tschudi, 1838 (Amphibia: Anura: Megophryidae) from Meghalaya, India. Records of the Zoological Survey of India 109: 91–108. https://doi.org/10.26515/rzsi/v109/i3/2009/158996
  • Matsui M (1997) Call characteristics of Malaysian Leptolalax with a description of two new species (Anura: Pelobatidae). Copeia volume: 158–165. https://doi.org/10.2307/1447851
  • Matsui M, Belabut DM, Ahmad N, Yong HS (2009) A new species of Leptolalax (Amphibia, Anura, Megophryidae) from Peninsular Malaysia. Zoological Science 26(3): 243–247. https://doi.org/10.2108/zsj.26.243
  • Matsui M, Panha S, Eto K (2023) Two new species of Leptobrachella from northern Thailand (Amphibia, Anura, Megophryidae). Current Herpetology Kyoto 42: 83–97. https://doi.org/10.5358/hsj.42.83
  • Matsui M, Zainudin R, Nishikawa K (2014b) A new species of Leptolalax from Sarawak, western Borneo (Anura: Megophryidae). Zoological Science 31(11): 773–779. https://doi.org/10.2108/zs140137
  • Nguyen LT, Poyarkov Jr NA, Le DT Vo BD, Ninh HT, Duong TV, Murphy RW, Sang NV (2018) A new species of Leptolalax (Anura: Megophryidae) from Son Tra Peninsula, central Vietnam. Zootaxa 4388(1): 1–21. https://doi.org/10.11646/zootaxa.4388.1.1
  • Nguyen LT, Tapley B, Nguyen CT, Luong HV, Rowley JJL (2021) A new species of Leptobrachella (Anura, Megophryidae) from Mount Pu Ta Leng, northwest Vietnam. Zootaxa 5016(3): 301–332. https://doi.org/10.11646/zootaxa.5016.3.1
  • Ninh HT, Nguyen TT, Le LTH, Nguyen TV, Quoc HN, Orlov NL, Bezman-Moseyko O, Le MV, Nguyen SN, Ziegler T (2024) A new Leaf-litter Frog of Leptobrachella (Anura: Megophryidae) from Tay Con Linh Mountain, northern Vietnam. Russian Journal of Herpetology 31(4): 191–224. https://doi.org/10.30906/1026-2296-2024-31-4-191-224
  • Ohler A, Marquis O, Swan S, Grosjean S (2000) Amphibian biodiversity of Hoang Lien Nature Reserve (Lao Cai Province, northern Vietnam) with description of two new species. Herpetozoa (Wien) 13(1/2): 71–87.
  • Ohler A, Wollenberg KC, Grosjean S, Hendrix R, Vences M, Ziegler T, Dubois A (2011) Sorting out Lalos: Description of new species and additional taxonomic data on megophryid frogs from northern Indochina (genus Leptolalax, Megophryidae, Anura). Zootaxa 3147(1): 1–83. https://doi.org/10.11646/zootaxa.3147.1.1
  • Pope CH (1931) Notes on Amphibians from Fukien, Hainan, and other parts of China. Bulletin of the AMNH, v. 61, article 8.
  • Poyarkov NJ, Rowley JJL, Gogoleva SI, Vassilieva AB, Galoyan EA, Orlov NL (2015) A new species of Leptolalax (Anura: Megophryidae) from the western Langbian Plateau, southern Vietnam. Zootaxa 3931(2): 221–252. https://doi.org/10.11646/zootaxa.3931.2.3
  • Qian TY, Xia X, Cao Y, Xiao NW, Yang DD (2020) A new species of the genus Leptobrachella (Anura: Megophryidae) Smith, 1925 from Wuling Mountains in Hunan Province, China. Zootaxa 4816(4): 491–526. https://doi.org/10.11646/zootaxa.4816.4.4
  • Rao DQ (2022) [2020] Atlas of Wildlife in Southwest China: Amphibian [in Chinese]. Zhu JG, Rao DQ eds., Atlas of Wildlife in Southwest China: Amphibian: 1–448 (printed in 2020 but not distributed until 2022). Beijing, Beijing Publishing Group.
  • Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choicem across a large model space. Systematic Biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029
  • Rowley JJL, Hoang DH, Le TTD, Dau QV, Cao TT (2010a) A new species of Leptolalax (Anura: Megophryidae) from Vietnam and further information on Leptolalax tuberosus. Zootaxa 2660: 33–45. https://doi.org/10.11646/zootaxa.2660.1.3
  • Rowley JJL, Stuart BL, Neang T, Hoang HD, Dau VQ, Nguyen TT, Emmett DA (2015a) A new species of Leptolalax (Anura: Megophryidae) from Vietnam and Cambodia. Zootaxa 4039(3): 401–417. https://doi.org/10.11646/zootaxa.4039.3.1
  • Rowley JJL, Tran DTA, Frankham GJ, Dekker AH, Le DTT Nguyen TQ, Dau VQ, Hoang HD (2015b) Undiagnosed cryptic diversity in small, microendemicfrogs (Leptolalax) from the central highlands of Vietnam. PLoS One 10(5): e0128382. https://doi.org/10.1371/journal.pone.0128382
  • Rowley JJL, Tran DTA, Le DTT Dau VQ, Peloso PLV, Nguyen TQ, Hoang HD, Nguyen TT, Ziegler T (2016) Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam. Zootaxa 4085(1): 63–102. https://doi.org/10.11646/zootaxa.4085.1.3
  • Rowley JJL, Dau VQ, Hoang HD, Le DTT Cutajar TP, Nguyen TT (2017a) A new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Zootaxa 4243(3): 544–564. https://doi.org/10.11646/zootaxa.4243.3.7
  • Sengupta S, Sailo S, Lalremsanga HT, Das A, Das I (2010) A new species of Leptolalax (Anura: Megophryidae) from Mizoram, north-eastern India. Zootaxa 2406(1): 56–68. https://doi.org/10.11646/zootaxa.2406.1.3
  • Shi SC, Hou YM, Song ZB, Jiang JP, Wang B (2021) A new leaf litter toad of Leptobrachella Smith, 1925 (Anura, Megophryidae) from Sichuan Province, China with supplementary description of L. oshanensis. Asian Herpetological Research 12(2): 143–166. https://doi.org/10.16373/j.cnki.ahr.200118
  • Shi SC, Shen T, Wang X, Jiang JP, Wang B (2023) Multiple data sources reveal a new Asian Leaf Litter Toad of Leptobrachella Smith, 1925 (Anura, Megophryidae) from southwestern China. Asian Herpetological Research 14(1): 65–94. https://doi.org/10.16373/j.cnki.ahr.220065
  • Smith MA (1925) Contributions to the herpetology of Borneo. Sarawak Museum Journal 3: 15–34.
  • Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG (1997) The CLUSTAL_X windows interface: Flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 25(24): 4876–4882. https://doi.org/10.1093/nar/25.24.4876
  • Wang J, Yang JH, Li Y, Lyu ZT, Zeng ZC, Liu ZY, Ye YH, Wang YY (2018) Morphology and molecular genetics reveal two new Leptobrachella species in southern China (Anura, Megophryidae). ZooKeys 776: 105–137. https://doi.org/10.3897/zookeys.776.22925
  • Wang J, Li YL, Li Y, Chen HH, Zeng YJ, Shen JM, Wang YY (2019) Morphology, molecular genetics, and acoustics reveal two new species of the genus Leptobrachella from northwestern Guizhou Province, China (Anura, Megophryidae). ZooKeys 848: 119–154. https://doi.org/10.3897/zookeys.848.29181
  • Wang J, Lyu ZT, Qi S, Zeng ZC, Zhang WX, Lu LS, Wang YY (2020) Two new Leptobrachella species (Anura, Megophryidae) from the Yunnan-Guizhou Plateau, southwestern China. ZooKeys 995: 97–125. https://doi.org/10.3897/zookeys.995.55939
  • Wang J, Qi S, Dai KY, Lyu ZT, Zeng ZC, Chen HH, Li YQ, Zhao YY, Wang YZ, Wang YY (2022) A new Leptobrachella species (Anura, Megophryidae) from South China, with comments on the taxonomic status of L. chishuiensis and L. purpurus. Zoosystematics and Evolution 98(1): 165–180. https://doi.org/10.3897/zse.98.73162
  • Wu YH, Yu ZB, Yang S, Duan ZP, Zuo AR, Zhang DC, Kilunda FK, Murphy RW, Che J (2025) Morphological and molecular evidence for a new species of the genus Leptobrachella (Anura, Megophryidae) from Gaoligong Mountain Range, Yunnan, China. Zoosystematics and Evolution 101(2): 449–463. https://doi.org/10.3897/zse.101.135560
  • Yu GD, Qin K, Meng T, Li P, Peng W, Chen WC (2024) A new species of the genus Leptobrachella (Amphibia, Anura, Megophryidae) from Dayaoshan National Nature Reserve, Guangxi, China. ZooKeys 1219: 105–122. https://doi.org/10.3897/zookeys.1219.121027
  • Yang JH, Wang YY, Chen GL, Rao DQ (2016) A new species of the genus Leptolalax (Anura: Megophryidae) from Mt. Gaoligongshan of western Yunnan Province, China. Zootaxa 4088: 379–394. https://doi.org/10.11646/zootaxa.4088.3.4
  • Yang JH, Zeng ZC, Wang YY (2018) Description of two new sympatric species of the genus Leptolalax (Anura: Megophryidae) from western Yunnan of China. PeerJ 6(e4586): 1–32. https://doi.org/10.7717/peerj.4586
  • Yuan ZY, Sun RD, Chen JM, Rowley JJL, Wu ZJ, Hou SB, Wang SN, Che J (2017) A new species of the genus Leptolalax (Anura: Megophryidae) from Guangxi, China. Zootaxa 4300(4): 551–570. https://doi.org/10.11646/zootaxa.4300.4.5

Supplementary material

Supplementary material 1 

Collection localities, voucher data, and Genbank accession numbers (16S rRNA) for all Leptobrachella samples used in this study.

Zhong Huang, Zhao-Chi Zeng, Jian Wang, Hao-Tian Wang, Zhen-Hai Deng, Zhi-Wei Wang, Xiao-Wen Liao, Yun-Ming Mo

Data type: docx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (28.70 kb)
login to comment