Zoosyst. Evol. 98 (1) 2022, 165-180 | DO! 10.3897/zse.98.73162 yee BERLIN A new Leptobrachella species (Anura, Megophryidae) from South China, with comments on the taxonomic status of L. chishuiensis and L. purpurus Jian Wang! *34, Shuo Qi!:*3, Ke-Yuan Dai®, Zhi-Tong Lyu!*%, Zhao-Chi Zeng!*>4, Hong-Hui Chen! *?4, Yuan-Qiu Li®, Yong-You Zhao°, Yun-Ze Wang®, Ying- Yong Wang?* State Key Laboratory of Biocontrol, Sun Yat-sen University, Guangzhou 510275, China The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, China School of Ecology, Sun Yat-sen University, Guangzhou 510275, China Shuanghuamu Biological Technology Co., Ltd, Shenzhen 518000, China Guangdong Shimentai National Nature Reserve, Oingyuan 513000, China Dn OF WDY KF Guangzhou Nanyang International School, Guangzhou 510620, China http://zoobank.org/EE3 14DBE-4650-4C50-B1D2-953F DD47F 876 Corresponding author: Ying- Yong Wang (wangyy@mail.sysu.edu.cn) Academic editor: Rafe Brown # Received 18 August 2021 # Accepted 16 May 2022 Published 3 June 2022 Abstract A new species of Leaf Litter Toad, Leptobrachella shimentaina sp. 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 charac- teristics 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 com- parisons of discrete character differences. In addition, we also discuss taxonomic uncertainty surrounding the identity of two congeners, L. chishuiensis and L. purpurus, which we interpret as indicative of taxonomic inflation in the species-rich subfamily Megophryidae. Key Words Leptobrachella shimentaina sp. nov., L. chishuiensis, L. purpurus, morphology, taxonomy Introduction The genus Leptobrachella Smith, 1925 recently was found to be paraphyletic with Leptolalax Dubois, 1983 based on a comprehensive molecular analysis combining fragments of mitochondrial and nuclear DNA markers by Chen et al. (2018). They suggested to synonymize two genera on account of their results clearly nested Lepto- brachella within Leptolalax, though the type species Leptobrachella mjobergi Smith, 1925 was not included. With 92 species, the forest-dependent genus Leptobrach- ella is widely distributed in southern China, Myanmar, northeastern India, Indochina region, Borneo and Natuna Island (Frost 2022). Species diversity in Leptobrachella may be underestimated, as suggested by 15 undescribed species proposed by Chen et al. (2018). One of the most widely-distributed species, Leptobra- chella liui (Fei & Ye, 1990) has been reported from Fujian, Guangdong, Guizhou, Hunan, and Zhejiang, provinces, Guangxi Zhuang Autonomous Region, and Hong Kong Copyright Wang, J. et al. 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. 166 SAR, China (Fei and Ye 1992; Fei et al. 2009; Fei et al. 2012). Later, Li et al. (2011) recognized Leptobrachel- la populations from Hong Kong and Guangdong (Xinyi City, Fengkai County, Shenzhen City) as the species L. pelodytoides (Boulenger, 1893). However, more recently, populations in Shenzhen of Guangdong and Hong Kong have been proposed as another species, L. /aui (Sung, Yang & Wang, 2014), and those in Xinyi have been de- scribed as L. yunkaiensis Wang, Li, Lyu & Wang, 2018. Although the taxonomic status of the population from Fengkai, western Guangdong remained unresolved due to the lack of molecular data, another noteworthy lineage from Shimentai Nature Reserve, northern Guangdong was indicated by Chen et al. (2018), as a putatively unde- scribed species (“Leptobrachella sp. 6”). In this paper we evaluate discrete character state differ- ences and phylogenetic relationships of seven additional specimens from Shimentai Nature Reserve and a single specimen from the adjacent Luokeng Nature Reserve (northern Guangdong, southern China Fig. 1), which sub- stantiate the recognition of “Leptobrachella sp. 6” (Chen et al. 2018) as a distinctive new species. Material and methods Phylogenetic analyses Eighteen new individuals were sequenced for phyloge- netic analyses, and 71 sequences were obtained from 4 42-43 fa 44-46 a 38—40 Guizhou Yunnan 36-51% } WGS 1984 Q® 100km i Wang, J. et al.: A new species of Leptobrachella GenBank (Suppl. material 1: Table S1). Our sampling includes individuals from most recognized congeners from China and neighboring countries. The 16S ribosom- al RNA mitochondrial gene (16S rRNA) fragment was sequenced for new samples; DNA extraction, PCR, and sequencing follow Wang et al. (2020). Sequences were aligned with Clustal X 2.0 (Thomp- son et al. 1997) with default parameters. For GenBank se- quences missing intervening sequence segments, we filled blank nucleotide positons “N” to indicate missing data. The aligned data was trimmed using default parameters and allowing no gap positions in Gblocks version 0.91b (Castresana 2000). 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 Revers- ible model, with variable sites modelled according to the Gamma distribution, and a portion of the sites invariant). Phylogenetic analysis was conducted using Bayesian in- ference (BI) in MrBayes 3.2.4 (Ronquist et al. 2012) and maximum likelihood (ML) in maximum likelihood (ML) in Randomized Axelerated Maximum Likelihood (RAx- ML, 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 Car- lo 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 Zhejiang aa @ 4 28 a 24 Taiwan ag E= \Hong Kong a wor Hainan Figure 1. Localities of Leptobrachella shimentaina sp. nov. (samples ID 1—10, Shimentai Nature Reserve, Guangdong, China; sample ID 11, Luokeng Nature Reserve, Guangdong, China); Leptobrachella bijie (samples ID 38—40, Zhaozishan Nature Reserve, Guizhou, China; sample ID 41, Chishui, Guizhou, China; samples ID 42-43, Huagaoxi Nature Reserve, Guizhou, China); Lepto- brachella chishuiensis (samples ID 44—46, Chishui, Guizhou, China); Leptobrachella alpina (samples ID 50-51, Mt Huangcaoling, Yunnan, China; sample ID 52, Pu’er, Yunnan, China); and Leptobrachella purpurus (samples ID 53-54, Yingjiang, Yunnan, China). Numbers correspond to the ID numbers in Suppl. material 1: Table S1. zse.pensoft.net Zoosyst. Evol. 98 (1) 2022, 165-180 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 hav- ing ML bootstrap values (BS) > 70 and BI posterior prob- abilities (BPP) > 0.95 were considered well supported. Morphology Our specimens of the putatively unnamed species were collected during opportunistic night searches. All speci- mens were euthanized, fixed in 5% buffered formalin for five hours, and then preserved in 70% ethanol, and sub- sequently deposited in The Museum of Biology, Sun Yat- sen University (SYS) and Chengdu Institute of Biology, Chinese Academy of Sciences (CIB), China. Following Fei et al. (2009) and Rowley et al. (2013), measurements were taken with digital calipers (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 167 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 diam- eter (EYE) from the anterior corner of the eye to poste- rior corner of the eye; internasal distance (IND); interor- bital 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 proxi- mal edge of the inner metatarsal tubercle; length of low- er 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 direct observation of calling in life, the presence of internal vocal sac openings (males), or the presence of eggs, seen via external inspection of the abdomen (females). Comparative morphological data for other congeneric species of Leptobrachella were ob- tained from museum specimens (Appendix 1) and from the literature (Table 1). Table 1. Data source of the currently known species of the genus Leptobrachella. ID Leptobrachella species 1 L. aerea (Rowley, Stuart, Richards, Phimmachak & Sivongxay, 2010c) 2 __L. aspera Wang, Lyu, Qi & Wang, 2020 3 _L. alpina (Fei, Ye & Li, 1990) 4 _ L. applebyi (Rowley & Cao, 2009) 5 _ L. arayai (Matsui, 1997) 6 __L. ardens (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) 7 L. baluensis Smith, 1931 8 __L. bashaensis Lyu, Dai, Wei, He, Yuan, Shi, Zhou, Ran, Kuang, Guo, Wei & Yuan, 2020 9 LL. bijie Wang, Li, Li, Chen & Wang, 2019 10 _L. bidoupensis (Rowley, Le, Tran & Hoang, 2011) 11 __L. bondangensis Eto, Matsui, Hamidy, Munir & Iskandar, 2018 12 _L. botsfordi (Rowley, Dau & Nguyen, 2013) 13 _L. bourreti (Dubois, 1983) 14 LL. brevicrus Dring, 1983 15___L. crocea (Rowley, Hoang, Le, Dau & Cao, 2010) 16 __L. chishuiensis Li, Liu, Wei & Wang, 2020 17 ___L. damingshanensis Chen, Yu, Cheng, Meng, Wei, Zhou & Lu, 2021 18 ___L. dorsospina Wang, Lyu, Qi & Wang, 2020 19 __L. dringi (Dubois, 1987) 20 iL. eos (Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011) 21 _L. feiiChen, Yuan & Che, 2020 22 _L. firthi (Rowley, Hoang, Dau, Le & Cao, 2012) 23 _L. fritinniens (Dehling & Matsui, 2013) 24 _L. fuliginosa (Matsui, 2006) 25 _L. flaviglandulosa Chen, Wang & Che, 2020 26 LL. fusca Eto, Matsui, Hamidy, Munir & Iskandar, 2018 27 _L. gracilis (Giinther, 1872) 28 _L. hamidi (Matsui, 1997) 29 LL. graminicola Nguyen, Tapley, Nguyen, Luong & Rowley, 2021 30 __L. heteropus (Boulenger, 1900) L. w = isos (Rowley, Stuart, Neang, Hoang, Dau, Nguyen & Emmett, 2015) . itiokai Eto, Matsui & Nishikawa, 2016 L. juliandringi Eto, Matsui & Nishikawa, 2015 L. jinshaensis Cheng, Shi, Li, Liu, Li & Wang, 2021 L. kajangensis (Grismer, Grismer & Youmans, 2004) WWW WwW W oO ol FP W DN — L. kalonensis (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) Literature obtained Rowley et al. 2010c Wang et al. 2020 Fei et al. 1990; Fei et al. 2009, 2016 Rowley and Cao 2009 Matsui 1997 Rowley et al. 2016 Dring 1983; Eto et al. 2016 Lyu et al. 2020a Wang et al. 2019 Rowley et al. 2011 Eto et al. 2018 Rowley et al. 2013 Ohler et al. 2011 Dring 1983; Eto et al. 2015 Rowley et al. 2010a Li et al. 2020 Chen et al. 2021b Wang et al. 2020 Inger et al. 1995; Matsui and Dehling 2012 Ohler et al. 2011 Chen et al. 2020 Rowley et al. 2012 Dehling and Matsui 2013 Matsui 2006 Chen et al. 2020 Eto et al. 2018 Gunther 1872; Dehling 201 2b Matsui 1997 Nguyen et al. 2021 Boulenger 1900 Rowley et al. 2015a Eto et al. 2016 Eto et al. 2015 Cheng et al. 2021 Grismer et al. 2004 Rowley et al. 2016 zse.pensoft.net 168 Wang, J. et al.: A new species of Leptobrachella ID Leptobrachella species Literature obtained 37 __L. kecil (Matsui, Belabut, Anmad & Yong, 2009) Matsui et al. 2009 38 _L. khasiorum (Das, Tron, Rangad & Hooroo, 2010) Das et al. 2010 39 _L. lateralis (Anderson, 1871) Anderson 1871; Humtsoe et al. 2008 40 LL. laui (Sung, Yang & Wang, 2014) Sung et al. 2014 41 LL. liui (Fei & Ye, 1990) Fei et al. 1990; Fei et al. 2009; Sung et al. 2014; this study 42 L. macrops (Duong, Do, Ngo, Nguyen & Poyarkov, 2018) Duong et al. 2018 43 _L. maculosa (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) Rowley et al. 2016 44 __L. mangshanensis (Hou, Zhang, Hu, Li, Shi, Chen, Mo & Wang, 2018) Hou et al. 2018; this study 45 LL. maoershanensis (Yuan, Sun, Chen, Rowley & Che, 2017) Yuan et al. 2017 46 _L. marmorata (Matsui, Zainudin & Nishikawa, 2014b) Matsui et al. 2014b 47 LL. maura (Inger, Lakim, Biun & Yambun, 1997) Inger et al. 1997 48 _L. melanoleuca (Matsui, 2006) Matsui 2006 49 _L. melica (Rowley, Stuart, Neang & Emmett, 2010) Rowley et al. 2010b 50 _L. minima (Taylor, 1962) Taylor 1962; Ohler et al. 2011 51 —_L. mjobergi Smith, 1925 Eto et al. 2015 52 L. murphyi Chen, Suwannapoom, Wu, Poyarkov, Xu, Pawangkhanant & Che, 2021 Chen et al. 2021a 53 _L. nahangensis (Lathrop, Murphy, Orlov & Ho, 1998) Lathrop et al. 1998 54 L. natunae (Gunther, 1895) Gunther 1895 55. namdongensis Hoang, Nguyen, Luu, Nguyen & Jiang, 2019 Hoang et al. 2019 56 _L. neangi Stuart & Rowley, 2020 Stuart and Rowley 2020 57 __L. niveimontis Chen, Poyarkov, Yuan & Che, 2020 Chen et al. 2020 58 —_L. nokrekensis (Mathew & Sen, 2010) Mathew and Sen 2010 59 __L. nyx (Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011) Ohler et al. 2011 60 _L. oshanensis (Liu, 1950) Liu, 1950; Shi et al. 2021 61 _L. pallida (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) Rowley et al. 2016 62 _L. palmata Inger & Stuebing, 1992 Inger and Stuebing 1992 63 _L. parva Dring, 1983 Dring 1983 64 _L. pelodytoides (Boulenger, 1893) Boulenger 1893; Ohler et al. 2011 65 ___L. petrops (Rowley, Dau, Hoang, Le, Cutajar & Nguyen, 2017) Rowley et al. 2017a 66 __L. picta (Malkmus, 1992) Malkmus 1992 67 _L. platycephala (Dehling, 2012) Dehling 2012a 68 __L. pluvialis (Ohler, Marquis, Swan & Grosjean, 2000) Ohler et al. 2000, 2011 69 _L. puhoatensis (Rowley, Dau & Cao, 2017) Rowley et al. 2017b 70 _L. purpurus (Yang, Zeng & Wang, 2018) Yang et al. 2018 71 __L. purpuraventra Wang, Li, Li, Chen & Wang, 2019 Wang et al. 2019 72 __L. pyrrhops (Poyarkov, Rowley, Gogoleva, Vassilieva, Galoyan & Orlov, 2015) Poyarkov et al. 2015 73 _L. rowleyae (Nguyen, Poyarkov, Le, Vo, Ninh, Duong, Murphy & Sang, 2018) Nguyen et al. 2018 74 _L. sabahmontana (Matsui, Nishikawa & Yambun, 2014) Matsui et al. 2014a 75 ___L. serasanae Dring, 1983 Dring 1983 76 _—__L. shangsiensis Chen, Liao, Zhou & Mo, 2019 Chen et al. 2019 77 _L. sola (Matsui, 2006) Matsui 2006 78 _L. sulyangensis Luo, Xiao, Gao & Zhou, 2020 Luo et al. 2020 79 __L. sungi (Lathrop, Murphy, Orlov & Ho, 1998) Lathrop et al. 1998 80 ___L. shiwandashanensis Chen, Peng, Pan, Liao, Liu & Huang, 2021 Chen et al. 2021c 81 ___L. tadungensis (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) Rowley et al. 2016 82 _L. tamdil (Sengupta, Sailo, Lalremsanga, Das & Das, 2010) Sengupta et al. 2010 83 _L. tengchongensis (Yang, Wang, Chen & Rao, 2016) Yang et al. 2016 84 LL. tuberosa (Inger, Orlov & Darevsky, 1999) Inger et al. 1999 85 ___L. ventripunctata (Fei, Ye & Li, 1990) Fei et al. 1990; Fei et al. 2009, 2016 86 _L. wuhuangmontis Wang, Yang & Wang, 2018 Wang et al. 2018 87 __L. wulingensis Qian, Xia, Cao, Xiao & Yang, 2020 Qian et al. 2020 89 _L. yingjiangensis (Yang, Zeng & Wang, 2018) Yang et al. 2018 90 LL. yunkaiensis Wang, Li, Lyu & Wang, 2018 Wang et al. 2018 91 _L. yeae Shi, Hou, Song, Jiang & Wang, 2021 Shi et al. 2021 92 __L. zhangyapingi (Jiang, Yan, Suwannapoom, Chomdej & Che, 201 3) Jiang et al. 2013 Results form a monophyletic clade, without structure of with- Essentially identical topologies were obtained by our ML and BI analyses (Fig. 2). In both phylogenetic estimates, all Leptobrachella samples formed a strongly-supported monophyletic clade. All Leptobrachella samples from populations from Shimentai and Luokeng nature reserves zse.pensoft.net in-population genetic divergence and with strong support (BPP = 1.00, BS = 100). The Shimentai and Luokeng lin- eage represents an independent lineage within Leptobra- chella, relatively close to the lineages of L. Jiui, L. mang- shanensis, L. maoershanensis, L. bashaensis, L. laui, and L. yunkaiensis. Nevertheless, support for the large clade Zoosyst. Evol. 98 (1) 2022, 165-180 18 23 19 20 21 is 77 | liui 1.00/99 1.42 13 14 1.00/86 16 24 roe h mangshanensis 1.007900} 22 6 28 8 6 7 10 @+4 | shimentaina sp. nov. 9 1 3 5 34 ; 100/100] “5°, | maoershanensis 35 1.00/100 3° | bashaensis 30 1.00/100 31 | laui 32 yunkaiensis 38 40 39 ny) bats 1.00/100| 143 bijie 44 chishuiensis 45 chishuiensis 46 chishuiensis 41 jinshaensis | ; suiyangensis niveimontis 1.00/99 47 (2H-48 purpuraventra 49 bourreti yeae 50 . 0.98/72-—1 51 | alpina | 0.99/83 52 53 1.00/94 {34 | purpurus orsospina 0.98/70 wulingensis tes tengchongensis (4) 1.00/100 namdongensis ————— puhoatensis (2) 1.00/99 petrops 1.00/79 khasiorum (3) 1.00/100 yingjiangensis flaviglandulosa @Lfe, >? | ventripunctata 1.00/99 aspera 0.02 a4 —<— ————— nahangensis nyx aerea He Ae vialis Leptobrachella eae = shangsiensis | wuhuangmontis SO. 1.00/92 sungi on zhangyapingi 0.97/89 botsfordi crocea 1.00/100 —________— Oreolalax rhodostigmatus Leptobrachium tengchongense 169 Figure 2. Bayesian inference tree derived from partial DNA sequences of the mitochondrial 16S r RNA gene. Numbers before slashes indicate Bayesian posterior probabilities and numbers after slashes are bootstrap support for maximum likelihood (1000 replicates) analyses. The number at the branch terminal corresponds to the ID in Suppl. material 1: Table S1. zse.pensoft.net 170 including these species is insignificant, therefore, the re- lationships among these species are still unresolved. De- tailed morphological examination has revealed discrete, diagnostic (non-overlapping ranges in traditional charac- ters) differences among the specimens from this unnamed lineage and all other congeners. Our phylogenetic result and morphological comparison unanimously agree with the proposal that the Shimentai and Luokeng lineage rep- resents an undescribed new species, i.e. “Leptobrachella sp. 6” in Chen et al. (2018). Therefore, based on the dis- crete and diagnostic morphological character differences, Leptobrachella populations from Shimentai and Luokeng nature reserves are described below as a new species. Taxonomic account Leptobrachella shimentaina J. Wang, Z.-T. Lyu & Y.- Y. Wang, sp. nov. http://zoobank.org/CF6EEF64-F539-415B-AD11-FA19CS56BC3D5 Shimentai Leaf Litter Toad (in English) / Shi Men Tai Zhang Tu Chan (41142 8in Chinese) Figs 3-6 Chresonymy. “Leptolalax sp.6” (Chen et al. 2018). Type material. Holotype 3. SYS a007684, from Jiu- zhou Yizhan (24°23'38.01"N, 113°06'35.38"E; ca. 300 m a.s.1.), Shimentai Nature Reserve, Yingde City, Guang- dong Province, China, collected by Jian Wang and Yong- You Zhao on 23 April 2019. Paratypes (n=7). 5 3: SYS a007683/CIB116079, same collection place as holotype; SYS a007685—7686, collected by Yu-Long Li and Hong-Hui Chen on 23 April 2019 from Shijiao (24°26'35.31"N, 113°09'21.55"E; ca 400 m a.s.l.), Shimentai Nature Reserve; SYS a007687, collected by Jian Wang and Yong-You Zhao on 24 April 2019 from Long- tankeng (24°26'31.38"N, 113°15'28.81"E; ca 310 m a.s.l.), Shimentai Nature Reserve; SYS a008329, collected by Jian Wang on 27 April 2020 from Luokeng Nature Reserve (24°31'42.3"N, 113°15'54.3"E; ca 590 m as.l.), Shaoguan City, Guangdong Province, China. 2 9: SYS a004711-4712, collected by Ying-Yong Wang, Jian Wang, and Zhi-Tong Lyu on 27 April 2016 from Hengshitang (24°26'14.92"N, 113°17'50.48"E; ca 380 ma.s.1.), Shimentai Nature Reserve. Etymology. The specific epithet “shimentaina” is an adjective derived from “shimentai’, referring to the type locality of the new species, Shimentai Nature Reserve, ‘ina’ 1s used as a feminine suffix which indicates the re- lationship of position. Diagnosis. (1) small body size [SVL 26.4—28.9 mm in six adult males, SVL 30.1 and 30.7 mm in two adult fe- males], (2) iris bicolored, upper half coppery orange and lower half silver, (3) tympanum distinct, (4) supratympan- ic line black, (5) fingers unwebbed, with distinct lateral fringes in males [absence in females], (6) toes with rudi- mentary webbing, lateral fringes wide in males [narrow in females], (7) longitudinal ridges under toes continuous, with constrictions at interphalangeal articulations, (8) zse.pensoft.net Wang, J. et al.: A new species of Leptobrachella heels slightly overlapping when adpressed, tibial-tarsal ar- ticulation reaching mid-orbit, (9) relative lengths of fingers I=l=IV IOD in L. chi- Shuiensis). The specimen SYS a004955 (sample ID 41 in our Figs 1, 2) from Chishui is clustered with the type specimens of L. chishuiensis (without appreciable genet- ic divergence), however, its morphological characters inversely match those of L. bijie. For instance, its heels are just meeting when thighs are positioned at right an- gles to the body (vs. heels overlapping in L. chishuiensis), the tibia-tarsal articulation reaches forward to the middle of the eye (vs. tympanum to eye in L. chishuiensis). Li et al. (2020) performed bioacoustic analysis to support proposed taxonomic validity of L. chishuiensis, howev- er, this study confused note and call intervals (comparing note interval in L. chishuiensis to call interval of L. bijie), which resulted in one apparently erroneous stated diag- nostic “difference” (a shorter call interval in L. chishui- ensis). The other difference (2 notes for L. bijie vs. 14 notes for L. chishuiensis) has been suggested to be only differences between populations rather than between spe- cies (Weaver et al. 2020). Geographically, the newly dis- covered population of L. bijie from southeastern Sichuan is much closer to the type locality of L. chishuiensis and any distinct barriers in geography (which might promote divergence) are absent. Therefore, we suspect L. chishui- ensis should most likely be considered conspecific (and be synonymized) with L. bijie. We also noticed the close relationship between the congeners Leptobrachella alpina and L. purpurus, \ead- ing to the problem of the taxonomic validity of L. purpu- rus. The latter species was described by Yang et al. (2018) based on two specimens, without comparison to the mo- lecular data of LZ. alpina. However, whereas molecular data corresponding to L. alpina were provided (Chen et al. 2018), corresponding data for L. purpurus were not in- cluded. In our phylogenetic analysis, both “species” clus- tered together, with minimal “interspecific” divergences, and with inconsistent support for their monophyly (BPP = 0.99, BS = 83; Fig. 2). Accessible voucher material for these two species is limited (two males for L. purpurus and three males for L. alpina), and reported type localities of the two species are distant from each other with a dis- tance over 340 km (Fig. 1). Therefore, we refrain from making taxonomic changes to L. purpurus until further studies, based on geopraphically comprehensive surveys, with robust, statistically-valid sample sizes, are conduct- ed throughout southwestern Yunnan Province, to clarify or refute species’ boundaries involving these two taxa. Zoosyst. Evol. 98 (1) 2022, 165-180 Acknowledgements We would like to thank Guangdong Shimentai Nature Re- serve, Guo-Xin Guo, Yang-Jin Zeng, Ming Kong, Ming-Ke Xu, Ming-Zhu Huang, Tai- Yang Zhou, Gui-Li Fan, Tian- Du Zhang, Di-Hao Wu, Run-Lin Li, and Hai-Long He, for their help with fieldwork, and Yao Li for her help in the lab work. We thank Rafe M. Brown, Kin Onn Chan, and the anonymous reviewer for their constructive comments on the manuscript. This work was supported by the Project of Fauna of Guangdong (DFGP202115), the National Animal Collection Resource Center, China, and the Project of An- imal Diversity Survey and Monitoring System Construc- tion of Guangdong Shimentai National Nature Reserve. References AmphibiaWeb (2022) AmphibiaWeb: information on amphibian biolo- gy and conservation, University of California, Berkeley, California. http://amphibiaweb.org/ [accessed 20 May 2022] Anderson J (1871) A list of the reptilian accession to the Indian Muse- um, 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. Boulenger GA (1900) Descriptions of new batrachians and reptiles from the Larut Hills, Perak. Annals & Magazine of Natural History 6(32): 186-194. https://doi.org/10.1080/00222930008678356 Castresana J (2000) Selection of conserved blocks from multiple align- ments for their use in phylogenetic analysis. Molecular Biology and Evolution 17(4): 540-552. https://doi.org/10.1093/oxfordjournals. molbev.a026334 Chen JM, Zhou WW, Poyarkov Jr NA, Stuart BL, Brown RM, Lathrop A, Wang YY, Yuan ZY, Jiang K, Hou M, Chen HM, Suwannapoom C, Nguyen SN, Duong TV, Papenfuss TJ, Murphy RW, Zhang YP, Che J (2017) A novel multilocus phylogenetic estimation reveals un- recognized diversity in Asian horned toads, genus Megophrys sensu lato (Anura: Megophryidae). Molecular Phylogenetics and Evolu- tion 106: 28-43. https://doi.org/10.1016/j.ympev.2016.09.004 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 diver- sity and historical biogeography of Asian leaf-litter frogs, genus Lepto- lalax (Anura: Megophryidae). Molecular Phylogenetics and Evolution 124: 162-171. https://doi.org/10.1016/j.ympev.2018.02.020 Chen WC, Liao X, Zhou SC, Mo YM (2019) A new species of Lepto- brachella (Anura: Megophryidae) from southern Guangxi, China. Zootaxa 4563(1): 67-82. https://doi.org/10.11646/zootaxa.4563.1.3 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 Re- search 41: 1-22. https://doi.org/10.24272/j.issn.2095-8137.2020.036 177 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, Yu GD, Cheng ZY, Meng T, Wei H, Zhou GY, Lu YW (2021b) A new species of Leptobrachella (Anura: Megophryidae) from central Guangxi, China. Zoological Research 42(6): 783-788. https://doi.org/10.24272/.issn.2095-8137.2021.179 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 spe- cies 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/zook- eys.1021.60729 Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: More models, new heuristics and parallel computing. Nature Methods 9(8): 772-772. https://doi.org/10.1038/nmeth.2109 Das I, Tron RKL, Rangad D, Hooroo RN (2010) A new species of Lep- tolalax (Anura: Megophryidae) from the sacred groves of Maw- phlang, Meghalaya, north-eastern India. Zootaxa 2339(1): 44-56. https://do1.org/10.11646/zootaxa.2339.1.2 Dehling JM (2012a) Eine neue Art der Gattung Leptolalax (Anura: Me- gophryidae) vom Gunung Benom, Westmalaysia/A new species of the genus Leptolalax (Anura: Megophryidae) from Gunung Benom, Peninsular Malaysia. Sauria 34: 9-21. Dehling JM (2012b) Redescription of Leptolalax gracilis (Gunther, 1872) from Borneo and taxonomic status of two populations of Lep- tolalax (Anura: Megophryidae) from Peninsular Malaysia. Zootaxa 3328(1): 20-34. https://do1.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. Dring J (1983) Frogs of the genus Leptobrachella (Pelobatidae). Amphib- ia-Reptilia 4(2): 89-102. https://doi.org/10.1163/156853883 X00012 Dubois A (1983) Note préliminaire sur le genre Leptolalax Dubois, 1980 (Amphibiens, Anoures), avec diagnose d’une espéce novelle du Vietnam. Alytes 2: 147-153. Dubois A (1987) “1986” Miscellanea taxinomica batrachologica (1). Alytes. Paris 5: 7-95. Duong TV, Do DT, Ngo CD, Nguyen TQ, Poyarkov Jr NA (2018) A new species of the genus Leptolalax (Anura: Megophryidae) from southern Vietnam. Zoological Research 39: 181-196. https://doi. org/10.24272/).issn.2095-8137.2018.009 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: Megophry- idae) from Kalimantan, Indonesia. Current Herpetology 37(2): 95- 105. https://doi.org/10.5358/hsj.37.95 zse.pensoft.net 178 Fei L, Ye CY (1992) Comments on the taxonomy of pelobatid toads of ge- nus 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. Publish- ing 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 & Technol- ogy, Chengdu, 619 pp. [In Chinese] Frost DR (2022) Amphibian Species of the World: an Online Reference. Version 6.1. Electronic Database. American Museum of Natural History, New York, USA. http://research.amnh.org/herpetology/am- phibia/index. html [accessed 20 May 2022] Grismer LL, Grismer JL, Youmans TM (2004) A new species of Lepto- lalax (Anura: Megophryidae) from Pulau Tioman, West Malaysia. Asiatic Herpetological Research 10: 8-11. Gunther A (1872) On the reptiles and amphibians of Borneo. Proceed- ings of the Scientific Meetings of the Zoological Society of London 1872: 586-600. Gunther 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. Singa- pore 67: 536-556. https://doi.org/10.26107/RBZ-2019-0042 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, Megophry- idae) from Hunan, China. Zootaxa 4444(3): 247-266. https://doi. org/10.11646/zootaxa.4444 3.2 Humtsoe LN, Bordoloi S, Ohler A, Dubois A (2008) Rediscovery of a long known species, /xalus lateralis Anderson, 1871. Zootaxa 1921(1): 24-34. https://doi.org/10.11646/zootaxa.1921.1.2 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 Lepto- lalax (Anura: Megophryidae) from Borneo. Asiatic Herpetological Research 7: 48-50. https://doi.org/10.5962/bhl part. 18855 Inger RF, Orlov N, Darevsky I (1999) Frogs of Vietnam: A report on new collections. Fieldiana. Zoology 92: 1-46. https://doi.org/10.5962/ bhl.title.3478 Jiang K, Yan F, Suwannapoom C, Chomde] S, Che J (2013) A new spe- cies 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 Lepto- lalax (Anura: Megophryidae) from northern Vietnam. Amphibia-Rep- tilia 19(3): 253-267. https://doi.org/10.1163/156853898X00160 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 zse.pensoft.net Wang, J. et al.: A new species of Leptobrachella Liu CC (1950) Amphibians of western China. Fieldiana. Zoology Mem- oires 2: 1-397. [+ 10 pl.] https://doi.org/10.5962/bhl.part.4737 Liu ZY, Chen GL, Zhu TQ, Zeng ZC, Lyu ZT, Wang J, Messenger K, Greenberg AJ, Guo ZX, Yang ZH, Shi SH, Wang YY (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 Luo T, Xiao N, Gao K, Zhou J (2020) A new species of Leptobrachella (Anura, Megophryidae) from Guizhou Province, China. ZooKeys 923: 115-140. https://doi.org/10.3897/zookeys.923.47172 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 Lyu ZT, Li YQ, Zeng ZC, Zhao J, Liu ZL, Guo GX, Wang YY (2020b) Four new species of Asian horned toads (Anura, Megophryidae, Megophrys) from southern China. ZooKeys 942: 105-140. https:// doi.org/10.3897/zookeys.942.47983 Lyu ZT, Zeng ZC, Wang J, Liu ZY, Huang YQ, Li WZ, Wang YY (2021) Four new species of Panophrys (Anura, Megophryidae) from east- ern China, with discussion on the recognition of Panophrys as a distinct genus. Zootaxa 4927(1): 009-040. https://doi.org/10.11646/ zootaxa.4927.1.2 Mahony S, Foley NM, Biju S, Teeling EC (2017) Evolutionary history of the Asian Horned Frogs (Megophryinae): Integrative approaches to timetree dating in the absence ofa fossil record. Molecular Biology and Evolution 34(3): 744-771. https://doi.org/10.1093/molbev/msw267 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 Lepto- brachium Tschudi, 1838 (Amphibia: Anura: Megophryidae) from Me- ghalaya, India. Records of the Zoological Survey of India 109: 91-108. Matsui M (1997) Call characteristics of Malaysian Leptolalax with a de- scription of two new species (Anura: Pelobatidae). Copeia volume: 158-165. https://doi.org/10.2307/1447851 Matsui M (2006) Three new species of Leptolalax from Thailand (Am- phibia, Anura, Megophryidae). Zoological Science 23(9): 821-830. https://doi.org/10.2108/zsj.23.821 Matsui M, Dehling JM (2012) Notes on an enigmatic Bornean mego- phryid, Leptolalax dringi Dubois, 1987 (Amphibia: Anura). Zootaxa 3317(1): 49-58. https://do1.org/10.11646/zootaxa.3317.1.4 Matsui M, Belabut DM, Ahmad N, Yong HS (2009) A new species of Lep- tolalax (Amphibia, Anura, Megophryidae) from Peninsular Malaysia. Zoological Science 26(3): 243-247. https://doi.org/10.2108/zs}.26.243 Matsui M, Nishikawa K, Yambun P (2014a) A new Leptolalax from the mountains of Sabah, Borneo (Amphibia, Anura, Megophryidae). Zoo- taxa 3753(3): 440-452. https://doi.org/10.11646/zootaxa.3753.5.3 Matsui M, Zainudin R, Nishikawa K (2014b) A New Species of Lepto- lalax from Sarawak, Western Borneo (Anura: Megophryidae). Zoo- logical Science 31(11): 773-779. https://doi.org/10.2108/zs 140137 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 Zoosyst. Evol. 98 (1) 2022, 165-180 Ohler A, Marquis O, Swan S, Grosjean S (2000) Amphibian biodiver- sity 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 Indo- china (genus Leptolalax, Megophryidae, Anura). Zootaxa 3147(1): 1-83. https://do1.org/10.11646/zootaxa.3147.1.1 Poyarkov NJ, Rowley JJL, Gogoleva SI, Vassilieva AB, Galoyan EA, Orlov NL (2015) A new species of Leptolalax (Anura: Me- gophryidae) from the western Langbian Plateau, southern Viet- nam. Zootaxa 3931(2): 221-252. https://doi.org/10.11646/zoot- axa.393. 2:3 Qi S, Lyu ZT, Wang J, Mo YM, Zeng ZC, Zeng YJ, Dai KY, Li YQ, Grismer LL, Wang YY (2021) Three new species of the genus Bou- lenophrys (Anura, Megophryidae) from southern China. Zootaxa 5072(5): 401-438. https://doi.org/10.11646/zootaxa.5072.5.1 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.48 16.4.4 Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Hohna 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, Cao TT (2009) A new species of Leptolalax (Anura: Me- gophryidae) from central Vietnam. Zootaxa 2198(1): 51-60. https:// doi.org/10.11646/zootaxa.2198.1.5 Rowley JJL, Hoang DH, Le TTD, Dau QV, Cao TT (2010a) A new spe- cies of Leptolalax (Anura: Megophryidae) from Vietnam and further information on Leptolalax tuberosus. Zootaxa 2660: 33-45. Rowley JJL, Stuart BL, Neang T, Emmett DA (2010b) A new species of Leptolalax (Anura: Megophryidae) from northeastern Cambodia. Zootaxa 2567(1): 57-68. https://doi.org/10.11646/zootaxa.2567.1.3 Rowley JJL, Stuart BL, Richards SJ, Phimmachak S, Sivongxay N (2010c) A new species of Leptolalax (Anura: Megophryidae) from Laos. Zootaxa 2681(1): 35-46. https://doi.org/10.11646/zoot- axa.2681.1.3 Rowley JJL, Le DTT, Tran DTA, Hoang DH (2011) A new species of Leptobrachella (Anura: Megophryidae) from southern Vietnam. Zootaxa 2796(1): 15-28. https://doi.org/10.11646/zootaxa.2796. 1.2 Rowley JJL, Hoang HD, Dau VQ, Le TTD, Cao TT (2012) A new spe- cies of Leptolalax (Anura: Megophryidae) from central Vietnam. Zootaxa 3321(1): 56—68. https://doi.org/10.11646/zootaxa.3321.1.4 Rowley JJL, Dau VQ, Nguyen TT (2013) A new species of Lepto- lalax (Anura: Megophryidae) from the highest mountain in Indo- china. Zootaxa 3737(4): 415-428. https://doi.org/10.11646/zoot- axa.3737.4.5 Rowley JJL, Stuart BL, Neang T, Hoang HD, Dau VQ, Nguyen TT, Emmett DA (2015a) A new species of Leptolalax (Anura: Mego- phryidae) from Vietnam and Cambodia. Zootaxa 4039(3): 401-417. https://do1.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 Diver- sity in Small, Microendemic Frogs (Leptolalax) from the Central Highlands of Vietnam. PLoS ONE 10(5): e0128382. https://doi. org/10.1371/journal.pone.0128382 179 Rowley JJL, Tran DTA, Le DTT, Dau VQ, Peloso PLV, Nguyen TQ, Hoang HD, Nguyen TT, Ziegler T (2016) Five new, microendem- ic Asian Leaf-litter Frogs (Leptolalax) from the southern Anna- mite 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 Rowley JJL, Dau VQ, Cao TT (2017b) A new species of Leptolalax (Anura: Megophryidae) from Vietnam. Zootaxa 4273(1): 61-79. https://do1.org/10.11646/zootaxa.4273.1.5 Sengupta S, Sailo S, Lalremsanga HT, Das A, Das I (2010) A new species of Leptolalax (Anura: Megophryidae) from Mizoram, north-east- ern India. Zootaxa 2406(1): 56-68. https://doi.org/10.11646/zoot- axa.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. os- hanensis. Asian Herpetological Research 12(2): 143-166. https:// doi.org/10.16373/j.cnki.ahr.200118 Silvestro D, Michalak I (2012) raxmlIGUI: A graphical front-end for RAXML. Organisms, Diversity & Evolution 12(4): 335-337. https:// doi.org/10.1007/s13127-011-0056-0 Smith MA (1925) Contributions to the herpetology of Borneo. Sarawak Museum Journal 3: 15-34. Stamatakis A (2006) RAxML-VI-HPC: Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics (Oxford, England) 22(21): 2688-2690. https://doi. org/10.1093/bioinformatics/btl446 Stuart BL, Rowley JJL (2020) A new Leptobrachella (Anura: Mego- phryidae) from the Cardamom Mountains of Cambodia. Zootaxa 4834(4): 556-572. https://doi.org/10.11646/zootaxa.4834.4.4 Sung YH, Yang JH, Wang YY (2014) A new species of Leptola- lax (Anura: Megophryidae) from southern China. Asian Her- petological Research 5(2): 80-90. https://doi.org/10.3724/ SP.J.1245.2014.00080 Taylor EH (1962) The amphibian fauna of Thailand. The University of Kansas Science Bulletin 43: 265-599. https://doi.org/10.5962/bhl. part. 13347 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. Nu- cleic Acids Research 25(24): 4876-4882. https://doi.org/10.1093/ nar/25.24.4876 Weaver S, Callaghan CT, Rowley JJL (2020) Anuran accents: Conti- nental-scale citizen science data reveal spatial and temporal patterns of call variability. Ecology and Evolution 10(21): 1-14. https://doi. org/10.1002/ece3.6833 Wane). Yane Jes ci byw Zio Zeng s Ze in ZyYieve.- 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://do1.org/10.3897/zookeys.848.29181 zse.pensoft.net 180 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 P, Liu Y, Huang W, Liu C (2021) An amphibian record of Si- chuan Province: Leptobrachella bijie. Sichuan Journal of Zo- ology 2021, 40(1): 66-69. https://doi.org/10.11984/).1ssn.1000- 7083.20200252 Yang JH, Wang YY, Chen GL, Rao DQ (2016) A new species of the genus Leptolalax (Anura: Megophryidae) from Mt. Gaoligongshan Appendix 1 Specimens examined: Leptobrachella alpina (n = 3): China: Yunnan: Jingdong County: Mt. Wuliang: CIB 24353-24354; SYS a 003927. Leptobrachella bijie (n = 9): China: Guizhou: Biyie City: SYS a007313—7320; China: Guizhou: Chishui City: SYS a004955. Leptobrachella laui (n = 26): China: Hong Kong: SYS a002057 (Holotype), SYS a002058; China: Guang- dong: Shenzhen City: SYSa 001505-—1507, 1515-— 1521, 3471-3472, 5644-5645. Leptobrachella liui (n = 32): China: Fujian: Mt. Wuy1: SYS a001572, 1596, 2478, 2479, 5925, 5926; Chi- na: Fujian: Mt. Datyun: SYS a001736, 6010; Chi- na: Fujian: Mt. Longqi: SYS a002505, 2506; China: Guangdong: Mt. Tongguzhang: SYS a004733-4735; China: Guangdong: Mt. Fenghuang: SYS a003698— 3699; China: Guangdong: Mt. Motianshi: SYS a007610—7613; China: Guangdong: Mt. Nankun: SYS a002020, 4497; China: Guangdong: Gutian Nature Reserve: SYS a002650; China: Jiangxi: Mt. Jiulian:; SYS a002104—2105; China: Jiangxi: Mt. Tongbo: SYS a001702, 2059; China: Jiangxi: Mt. Yangjifeng: SYS a006667, 6672; China: Zhejiang: Jingning: SYS a002732-—2735. Leptobrachella mangshanensis (n = 11): China: Hu- nan: Mangshan Nature Reserve: SYS a008366; China: Guangdong: Nanling Nature Reserve: SYS a002828—2830, 5754; China: Guangdong: Shimentai Nature Reserve: SYS a005763, 6880; China: Guang- dong: Mt. Tianjing: SYS a002806, 2809; China: Guangdong: Mt. Dadong: SYS a002847-2848. Leptobrachella purpuraventra (n= 15): China: Guizhou: Biyie City: SYS a007277—7284, 7300-7306. Leptobrachella yunkaiensis (n = 8): China: Guangdong: Maoming City: Dawuling Forestry Station: SYS a004664/CIB107272, SYS a004663, 4665-4669, 4690. zse.pensoft.net Wang, J. et al.: A new species of Leptobrachella 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: Mego- phryidae) from Guangxi, China. Zootaxa 4300(4): 551-570. https:// doi.org/10.11646/zootaxa.4300.4.5 Supplementary material | Table SI Authors: Jian Wang, Shuo Qi, Ke-Yuan Dai, Zhi-Tong Lyu, Zhao-Chi Zeng, Hong-Hui Chen, Yuan-Qiu L1, Yong- You Zhao, Yun-Ze Wang, Ying- Yong Wang Data type: excel file Explanation note: Collection localities, voucher data, and Genbank numbers (16S rRNA) for all Leptobrachella samples used in this study. Copyright notice: 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 us- ers 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. Link: https://doi.org/10.3897/zse.98.73162.suppl1 Supplementary material 2 Table S2 Authors: Jian Wang, Shuo Qi, Ke-Yuan Dai, Zhi-Tong Lyu, Zhao-Chi Zeng, Hong-Hui Chen, Yuan-Qiu L1, Yong-You Zhao, Yun-Ze Wang, Ying- Yong Wang Data type: excel file Explanation note: Uncorrected P-distance of 16S gene among 45 Leptobrachella species in this study. Copyright notice: 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 us- ers 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. Link: https://doi.org/10.3897/zse.98.73162.suppl2