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

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

We report the first country record of Leptobrachella eos (Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011) from Thailand, based on one specimen collected from Chom poo Phuka nature trail, Bo Kluea District. Morphologically, the specimen displayed good agreement with the original descriptions provided for L. eos. Phylogenetically, the specimens clustered according to the sequences of type locality of L. eos. Notably, our discovery increases the number of Leptobrachella species known to occur in Thailand to nine.

A new species of the Asian leaf litter toad genus Leptobrachella from central south China is described. Molecular phylogenetic analyses, based on mitochondrial 16S rRNA and nuclear RAG1 gene sequences indicated the new species as an independent clade in the genus. The new species could be distinguished from its congeners by a combination of the following characters: body of medium size (SVL 29.2–34.2 mm in 15 adult males and 34.4–43.1 mm in seven adult females); distinct black spots present on flanks; toes rudimentary webbed, with wide lateral fringes; ventral belly white with distinct nebulous brown speckling on ventrolateral flanks; skin on dorsum shagreened with fine tiny granules or short ridges; iris copper above, silver below; heels overlapped when thighs are positioned at right angles to the body; tibia-tarsal articulation reaches the middle eye; dorsal surface of tadpole semi-transparent light brown, spots on tail absent, keratodont row formula I: 3+3/2+2: I; call series basically consist of repeated long calls, at dominant frequency (5093 ± 412 Hz).

A new species of the genus Leptobrachella, Leptobrachella damingshanensis sp. nov. is described from central Guangxi, China, based on morphological, molecular, and acoustic data. Phylogenetic analyses using a 16S rRNA gene fragment indicated that the new species formed an independent clade close to L. nahangensis and L. nyx. However, the new species differs from all known congeners based on a combination of the following characters: medium size (snout-vent length (SVL) 33.6–34.4 mm in males); pair of distinct jacinth humeral glands on proximal thigh area; rough dorsal skin with sparse jacinth tubercles and several short longitudinal ridges; creamy white ventral surface with small, creamy white glands on throat, chest, belly, and ventral surfaces of thighs, becoming more concentrated near lateral margin; webbing and lateral fringes absent on fingers, toes with rudimentary webbing and narrow lateral fringes; flanks with small irregular black spots; iris bicolored, upper half copper, fading to silver in lower half. Uncorrected p-distances were greater than 4.7% between Leptobrachella damingshanensis sp. nov. and all homologous DNA sequences available for the 16S rRNA gene. The new species is only known to occur in the Damingshan National Nature Reserve, where it inhabits montane evergreen forests between 1 000–1 200 m a.s.l.

Based on a new herpetological collection from the Mu Cang Chai Species and Habitat Conservation Area, Yen Bai Province, we report two new records of amphibians, viz. Leptobrachella niveimontis Chen, Poyarkov, Yuan et Che, 2020 and Leptobrachella yingjiangensis (Yang, Zeng et Wang, 2018) from Vietnam. Morphological descriptions and ecological notes of afore mentioned species are provided on the basis of new materials. Our findings increase the species number of the genus Leptobrachella to 29 in Vietnam.

The Hoang Lien Range in northwest Vietnam is known to harbour five species of Leptobrachella, a specious genus of terrestrial frogs. We collected specimens of Leptobrachella from Mount Pu Ta Leng on the second highest peak in the Hoang Lien Range and use morphological and molecular data to show that this population represents a previously undescribed species which we name Leptobrachella graminicola sp. nov. after the abundance of calling males on sedge-like plants. This new species is closely related to L. bourreti but the new species is distinguished from L. bourreti and other congeneric species by a combination of the following characters: (1) a body size range of 23.1–24.6 mm in six adult males and 28.6–32.9 mm in five adult females; (2) skin on dorsum smooth, with many tubercles and lacking dermal ridges; (3) toes with rudimentary webbing and broad lateral fringes; (4) belly white with brown spots; (5) throat dark brown with light grey-brown flecks and spots; (6) a row of large white spots on the outer edge of the tarsus extending from the heel to the inner metatarsal tubercle, sometimes forming a long white stripe; and (7) a bicoloured iris. The new, high-elevation species is likely to be range-restricted and threatened by habitat loss and the activities of tourists.

Leptobrachella petrops (Rowley, Dau, Hoang, Le, Cutajar et Nguyen, 2017) was recently described from northern Vietnam. The tadpoles of this endemic species were collected for the first time from its type locality in Cham Chu Nature Reserve, Tuyen Quang Province. Taxonomic identification was based on a fragment of 16S rRNA gene of the newly collected tadpole and sequences of the type series obtained from GenBank. Morphological characters of the tadpoles (Gosner stages 25, 28, 35, 36, 43, 44, and 45) are as follows: Size medium, body elliptical, elongated, eyes are dorsolateral, narial apertures with characteristic gently rounded rims, slightly lifted above body surface, breathing hole on the left, vent tube dextral, separated from body and attached to lower fin, tail length exceeding two times body length, both upper and lower tail fins reach maximum height in posterior third, tip of tail pointed; dorsal surface olive-brown with distinct whitish-golden specklings, the tail has large black spots, ventral surface opaque-white, internal gills distinctly reddish. LTRF: 4(2 – 4)/3(1 – 2). In addition, we compare the tadpoles of Leptobrachella petrops with those of other species in the genus Leptobrachella.

We present a new state record of Leptobrachella tamdil from Manipur State, northeastern India based on three individuals collected from Chakpi stream, Chandel District. This record represents the range extension of the species as well as the easternmost distribution record. We also provide additional morphological data as well as the first time genetic data for the species and inferred its phylogenetic position using mitochondrial 16S rRNA marker gene sequence.

A new species of the Asian leaf litter toad genusLeptobrachellais described from Guizhou Province, China. Molecular phylogenetic analyses support the new species as an independent lineage deeply nested in theLeptobrachellaclade. The new species is distinguished from its congeners by a combination of the following morphological characters: body size medium (SVL 29.7–31.2 mm in five adult males); dorsal skin shagreened, some of the granules forming longitudinal short skin ridges; tympanum distinctly discernible, slightly concave; supra-axillary, femoral, pectoral and ventrolateral glands distinctly visible; absence of webbing and lateral fringes on fingers; toes with narrow lateral fringes but without webbing; heels overlapping when thighs are positioned at right angles to the body; tibia-tarsal articulation reaching the middle of eye when leg stretched forward. The discovery highlighted the underestimated species diversity in theLeptobrachellatoads in southwestern China.

A new species of the genus Leptobrachella, Leptobrachella damingshanensis sp. nov. is described from central Guangxi, China, based on morphological, molecular, and acoustic data. Phylogenetic analyses using a 16S rRNA gene fragment indicated that the new species formed an independent clade close to L. nahangensis and L. nyx. However, the new species differs from all known congeners based on a combination of the following characters: medium size (snout-vent length (SVL) 33.6–34.4 mm in males); pair of distinct jacinth humeral glands on proximal thigh area; rough dorsal skin with sparse jacinth tubercles and several short longitudinal ridges; creamy white ventral surface with small, creamy white glands on throat, chest, belly, and ventral surfaces of thighs, becoming more concentrated near lateral margin; webbing and lateral fringes absent on fingers, toes with rudimentary webbing and narrow lateral fringes; flanks with small irregular black spots; iris bicolored, upper half copper, fading to silver in lower half. Uncorrected p-distances were greater than 4.7% between Leptobrachella damingshanensis sp. nov. and all homologous DNA sequences available for the 16S rRNA gene. The new species is only known to occur in the Damingshan National Nature Reserve, where it inhabits montane evergreen forests between 1 000–1 200 m.

A new species of the Asian leaf litter toad genusLeptobrachellafrom Guizhou Province, China is described based on molecular phylogenetic analyses, morphological comparisons, and bioacoustics data. Phylogenetic analyses based on the mitochondrial 16S rRNA gene sequences supported the new species as an independent clade nested into theLeptobrachellaclade and sister toL. bijie. The new species could be distinguished from its congeners by a combination of the following characters: small body size (SVL 30.8–33.4 mm in seven adult males, and 34.2 mm in one adult female); dorsal skin shagreened, some of the granules forming longitudinal short skin ridges; tympanum distinctly discernible, slightly concave; internasal distance longer than interorbital distance; supra-axillary, femoral, pectoral and ventrolateral glands distinctly visible; absence of webbing and lateral fringes on fingers; toes with rudimentary webbing and shallow lateral fringes, relative finger lengths II < IV < I < III; heels overlapped when thighs are positioned at right angles to the body; and tibia-tarsal articulation reaches the tympanum.

Asian leaf-litter toads of the genus Leptobrachella represent a great anuran diversification in Asia. Previous studies have suggested that the diversity of this genus is still underestimated. During herpetological surveys from 2013 to 2018, a series of Leptobrachella specimens were collected from the international border areas in the southern and western parts of Yunnan Province, China. Subsequent analyses based on morphological and molecular data revealed three distinct and previously unknown lineages, which we formally describe as three new species herein. Among them, we describe a new species that occurs at the highest known elevation for Leptobrachella in China. Four species of Leptobrachella, including two new species, are found in the same reserve. Furthermore, our results suggest that the population from Longchuan County, Yunnan, may represent an additional new species of Leptobrachella, although we tentatively assigned it to Leptobrachella cf. yingjiangensis due to the small sample size examined. Lastly, we provide the first description of females of L. yingjiangensis. Our results further highlight that both micro-endemism and sympatric distributions of species are common patterns in Leptobrachella, that contribute to taxonomic and conservation challenges in these frogs. We provide an identification key for Leptobrachella known to occur in Yunnan. Given the lack of knowledge on species diversity of Leptobrachella along international border areas, we recommend that future studies include trans-boundary collaborative surveys.

Vassilieva, Anna B. (2021): Larval external morphology and chondrocranium of Leptobrachella bidoupensis (Anura: Megophryidae), an endemic litter toad from southern Vietnam. Zootaxa 5068 (4): 587-594, DOI: https://doi.org/10.11646/zootaxa.5068.4.8

Thailand is considered a global biodiversity hotspot that is known to harbour a striking diversity of endemic species. However, several research studies have determined that the level of amphibian diversity in the country has been significantly underestimated. The megophryid genus Leptobrachella Smith, 1925 is currently known to include 89 species that are primarily distributed throughout southern China and Southeast Asia; however, only seven species have been found in Thailand. Based on an integrative approach encompassing genetic and morphological analyses, we have concluded that the population identified from Chiang Rai Province of Thailand is conspecific with Leptobrachella ventripunctata (Fei, Ye, and Li, 1990). Importantly, this is the first confirmation record of this species, based on molecular and morphological evidence in Thailand. The discovery of this species reaffirms that the diversity within the genus has been underestimated with many species yet to be discovered. In addition, the findings of our study further highlight the lack of existing knowledge on amphibian taxonomy and an underestimation of the biodiversity that exists along these national border areas.

Leptobrachella murphyi sp. nov. Chen, Suwannapoom, Wu, Poyarkov, Xu, Pawangkhanant & Che (Figures 4–6) Chresonymy: Leptolalax sp. — Ohler et al. 2011: 9; Leptolalax sp. 3 — Chen et al. 2018: 165; Leptolalax pelodytoides — Taylor 1962. Holotype. KIZ034039, adult male, collected from Ban Huai Wok in Doi Inthanon, Chiang Mai Province, Thailand (98.5348°N, 18.4662°E, 587m a.s.l.; Fig. 1), on 17 January 2018 by Chatmongkon Suwannapoom. Paratype. KIZ034038, KIZ034165–66, KIZ034158, four adult females and KIZ034159, one adult male from the same locality as the holotype, collected by Chatmongkon Suwannapoom and Yunhe Wu on 17 January 2018; KIZ034182–83, two adult males, from Ban Khun Klang in Doi Inthanon, Chiang Mai Province, Thailand (98.5140°N, 18.6670°E, 1090 m a.s.l.; Fig. 1) collected by Yunhe Wu on 19 January 2018; KIZ031168, KIZ031198 – KIZ031200, four adult males, from Mae Ya Noi in Doi Inthanon, Chiang Mai Province, Thailand (98.5337°N, 18.4939°E, 956 m a.s.l.) collected by Chatmongkon Suwannapoom on 6 August 2018; KIZ034225, one adult male from Ban Khun Wang in Doi Inthanon, Chiang Mai Province, Thailand (98.5049°N, 18.6045°E) collected by Chatmongkon Suwannapoom on 19 January 2018; KIZ034173, one adult male from Pha Dok Siaw Waterfall in Doi Inthanon, Chiang Mai Province, Thailand (98.5305°N, 18.5410°E, 1200 m a.s.l.) collected by Yunhe Wu on 20 January 2018. Diagnosis. Leptobrachella murphyi sp. nov. can be distinguished from its congeners by the following combination of morphological characters: (1) body size small (SVL 23.2–24.9 mm in 10 adult males, 29.3–32.1 mm in 4 adult females); (2) tympanum distinct, almost entirely black; (3) skin on dorsum shagreened with reddish tubercles and folds; (4) creamy white belly with small black spots on the margin; (5) distinct reverse-triangle marking between eyes, close to a black W-shaped marking between axillae on the dorsum; (6) distinct black blotches on flanks; (7) white ventrolateral glands forming a distinct line; (8) finger webbing and fringes absent; (9) toe webbing rudimentary and lateral fringes wide; (10) longitudinal ridges distinct under toes and uninterrupted at the articulations; (11) iris distinctly bicolored, typically orange in upper half and silver white in lower half; (12) small pectoral gland embellished on the flesh-colored oval marking; and (13) supratympanic line distinct with reddish pigmentation. Description of the holotype. Adult male, SVL 24.60 mm; head length almost equal with width (HDL/ HDW=1.03), head triangular in dorsal view; snout rounded in both ventral view and lateral view, protruding slightly beyond lower jaw (Fig. 4); oval-shaped nostril closer to tip of snout than to anterior margin of eye; loreal region oblique; canthus rostralis indistinct; eyes large (EYE/HDL=0.37), eye diameter slightly shorter than snout length (EYE/SNT=0.97), eyes notably protuberant in both dorsal and lateral views, pupil vertical; tympanum distinct, rounded, tympanum diameter smaller than eye (TMP/EYE = 0.57); tympanic annulus notably elevated; vomerine teeth absent; vocal sac openings small, slit-like, located postero-laterally on mouth floor; supratympanic ridge distinct with reddish pigmentation, running from posterior corner of eye towards axilla, posterior end of supratympanic ridge not expanded (Fig. 4). Forelimb relatively long (FAL/SVL=0.26), fingers long and slender (ML/SVL=0.28), without webbing and lateral fringes (Fig. 4); relative length of fingers: IIColoration of holotype. In life, dorsal surface of head and trunk olive brown, with a distinct reverse-triangle dark markings between eyes connecting to a thick dark W-shaped marking between axillae; elbow to upper arm and tibio-tarsal articulation distinctly yellowish orange in color on the dorsum; transverse black bars present on dorsal surface of fingers and toes, lower arms, tarsus, thighs and tibia (Fig. 4); one dark blotch between nostril and eyes on loreal region; upper lip barred with black and white; supratympanic ridge reddish; a large black marking under supratympanic ridge covers most of tympanum; distinct, irregularly-shaped black blotches present on flanks from the groin to axilla; fine, distinct reddish tubercles scattered on upper eyelids, snout, dorsal surfaces of head, body and limbs. Ventral surface of throat pale with some dusting; ventral surface of belly creamy white with small spots on the margin; ventral surfaces of limbs pinkish, scattered with small white speckles. Ventrolateral glands, pectoral glands and femoral glands yellowish white. Iris distinctly bicolored, typically orange in upper half, silver white in lower half, with black reticulations throughout (Fig. 4). In preservation, dorsal surface of head and trunk dark brown with light tawny limbs. Ventral surface of throat, chest, belly and limbs creamy white; brown dusting present on ventral surfaces of throat. The patterns of dark markings, spots and bars all over the body are same as when in life. Bars and blotches on dorsum and limbs became more distinct. Pectoral glands indistinct; femoral glands distinct and white (Fig. 5). Morphological variation. All paratypes match the overall characters of the holotype (Table 3). Representative photographs of paratypes in life are shown in Fig. 6 and in preservation are shown in Fig. 5. Females larger than males (SVL 23.2–24.9 mm in 10 adult males and 29.3–32.1 mm in 4 adult females). All males have small slit-like vocal sac openings, as well as a comparatively large single subgular vocal sac when inflated during calling. Dusting pattern on throat and margin of belly varies individually (Fig. 6). Males have more distinct white flecks dorsum than females (Fig. 6). Etymology. The specific epithet “ murphyi ” is a patronymic noun in the genitive singular; derived from the name of Prof. Robert W. Murphy from the Royal Ontario Museum, Canada, the co-founder of ColdCode, the international effort to DNA barcode species of amphibians and reptiles. We acknowledge his continuous support and inspiration of our study across the country border from China to Southeast Asia. For the common name, we recommend “Chiang Mai Leaf Litter Toad”. Distribution and ecology. Leptobrachella murphyi sp. nov. is currently known from two isolated montane areas: Doi Inthanon and Doi Chiang Dao, both belonging to the Thanon Thong Chai Range in Chiang Mai Province, Thailand (Fig. 1). These two sites are separated by a straight-line distance of about 110 kilometers. The known localities in Doi Inthanon for this new species includes Mae Ya Noi, Ban Huai Wok, Ban Khun Klang, Khun Wang 1, Pha Dok Siaw Waterfall and Mae Ya at elevations ranging from 587 to 1477 m a.s.l. During our field surveys, males were usually located at night by their calling while sitting on the litter and leaves of bushes along the streams and waterfalls surrounded by moist evergreen broadleaved forests (Fig. 7). A tadpole of Leptobrachella, which was previously regarded as Leptobrachella sp. (GenBank No. JN848455; Ohler et al., 2011) and collected from Doi Chiang Dao, belongs to the new species we describe (Fig. 2). The ecological information on L. murphyi in Doi Chiang Dao is currently insufficient. Advertisement calls of the new species, resembling calling of orthopterans, could be heard along the streams from November to February in Doi Inthanon; a gravid female (KIZ034165) collected in January was swollen with pure white eggs. Comparisons. Compared with the 26 known congeners occurring south of the Isthmus of Kra, Leptobrachella murphyi sp. nov. can be easily 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 by the presence of supra-axillary and ventrolateral glands (vs. absence); and from the following miniaturized species it can be distinguished by presence of rounded digit tips (vs. pointed digit tips in all species listed below) and by having an obviously larger body size (SVL 23.2–24.9 mm in males of L. murphyi sp. nov.): L. baluensis (14.9–15.9 mm in males), L. brevicrus (17.1–17.8 mm in males), L. bondangensis (17.8 mm in males), L. fusca (16.3 mm in males), L. itiokai (15.2–16.7 mm in males), L. juliandringi (17.0– 17.2 mm in males), L. mjobergi (15.7–19.0 mm in males), L. natunae (17.6 mm in males), L. parva (15.0– 16.9 mm in males), L. palmata (14.4–16.8 mm in males) and L. serasanae (16.9 mm in female). Leptobrachella murphyi sp. nov. differs from all other species of Leptobrachella occurring north of the Isthmus of Kra by a combination of small body size, distinct black tympanum, reddish supratympanic line, skin texture and markings on dorsum, ventral coloration, white ventrolateral glands forming a distinct line, degree of webbing and lateral fringes on the toes, distinct black blotches on flanks, longitudinal ridges distinct under toes and uninterrupted at the articulations, small pectoral gland and a bicolored iris (Table 4). In particular, L. murphyi sp. nov. can be easily distinguished from its phylogenetically close congeners. It differs from L. oshanensis, by having relatively smaller body size in males (SVL 23.2–24.9 mm vs. 26.6–30.7 mm), femoral glands located closer to the knee than to the vent (vs. closer to the vent than to the knee), skin on dorsum shagreened with reddish tubercles and folds (vs. smooth with few glandular ridges), as well as the presence of webbing and dermal fringes on toes (vs. absence). Leptobrachella murphyi sp. nov. differs from L. tengchongensis by having a bicolored iris (vs. uniform), relatively larger body size in females (SVL 29.3–32.1 vs. 28.8–28.9), wide dermal fringes on toes (vs. narrow), ventrolateral glands distinct (vs. indistinct) and ventral side creamy white with small black spots on the margin (vs. ventral surfaces white with dark brown blotches). The new species differs from L. purpurus by having olive brown dorsum coloration in life (vs. purplish brown), creamy white belly with small black spots on the margin (vs. ventral side dull white with indistinct grey dusting), pectoral gland indistinct (vs. distinct), as well as the absence of black marking/spots on dorsum and flanks mottled with distinct yellow pigmentation al. 2017; Yuan et al. 2017; Yang et al. 2018; Chen et al. 2020). ......continued on the next page ......continued on the next page ......continued on the next page (vs. presence). Leptobrachella murphyi sp. nov. differs from L. purpuraventra by having relatively smaller body size (SVL 23.2–24.9 mm in males, 29.3–32.1 mm in females vs. 27.3–29.8 mm in males, 33.0– 35.3 mm in females), toes with wide lateral fringes (vs. narrow), ventral surface creamy white (vs. grey purple with distinct nebulous greyish speckling on chest and ventrolateral flanks), as well as the absence of dense, tiny conical spines on surface of chest extending to anterior region of abdomen in males (vs. presence). The new species can be distinguished from L. alpina by dorsum olive brown in life (vs. grey brown), indistinct pectoral gland (vs. distinct) and supra-axillary gland yellowish (vs. white). The new species differs from L. bijie by having smaller body size in males (SVL 23.2–24.9 mm vs. 29.0– 30.4 mm), toes with wide lateral fringes (vs. narrow), as well as the absence of distinct nebulous greyish speckling on chest and ventrolateral flanks (vs. presence) and the absence of dense tiny conical spines present on surface of chest in males during breeding season (vs. presence). Leptobrachella murphyi sp. nov. differs from L. eos by having a smaller body size in males (SVL 23.2–24.9 mm vs. 33.1–34.7 mm), olive brown dorsum with black markings and spots in life (vs. almost uniformly brown dorsal coloration), as well as the presence of distinct black spots on flanks (vs. absence) and the presence of distinct supratympanic fold (vs. absence). The new species differs from L. bourreti by having a relatively smaller body size in males (SVL 23.2–24.9 mm vs. 28.0– 36.2 mm), wide dermal fringes on toes (vs. narrow), skin on dorsum shagreened with reddish tubercles and folds (vs. relatively smooth, some with small warts). The new species differs from L. chishuiensis by having smaller body size in males (SVL 23.2–24.9 mm vs. 30.8–33.4 mm). The new species differs from L. dorsospina by having smaller body size in males (SVL 23.2–24.9 mm vs. 28.7–30.5 mm), toes with wide lateral fringes (vs. narrow), as well as the absence of dense conical spines on dorsum (vs. presence). The new species differs from L. jinshaensis by having smaller body size in males (SVL 23.2–24.9 mm vs. 29.7–31.2 mm) and toes with wide lateral fringes (vs. narrow). The new species differs from L. niveimontis by having toes with wide lateral fringes (vs. narrow) and belly creamy white (vs. belly marbled with black speckling). The new species differs from L. suiyangensis by having dermal ridges under toes not interrupted at articulations (vs. interrupted) and smaller body size in males (SVL 23.2–24.9 mm vs. 28.7–29.7 mm). The new species differs from L. wulingensis by having toes with wide lateral fringes (vs. narrow) and dermal ridges under toes not interrupted at articulations (vs. interrupted). The new species differs from L. yeae by having dermal ridges under toes not interrupted at articulations (vs. interrupted) and toes with wide lateral fringes (vs. narrow). For the remaining known species of Thai Leptobrachella, L. murphyi sp. nov. differs from L. fuliginosa by having a relatively smaller body size in males (SVL 23.2–24.9 mm vs. 28.2–30.0 mm), wide dermal fringes on toes (vs. narrow), distinct femoral glands (vs. narrow), skin on dorsum shagreened with reddish tubercles and folds (vs. skin nearly smooth, with a few tubercles), as well as the absence of dusty ventral side (vs. presence). The new species differs from L. melanoleuca by a relatively smaller body size (SVL 23.2–24.9 mm in males, 29.3–32.1 mm in females vs. 26.6–28.8 mm in males, 32.7 mm in females), creamy white belly with small black spots on the margin (vs. ventral coloration with large black markings on a white background), distinct femoral gland (vs. indistinct) and distinct ventrolateral glandular ridges (vs. indistinct). The new species differs from L. sola by having iris with orange in upper half and silver white in lower half (vs. iris with dark red in upper half and dark brown in lower half), as well as the presence of supra-axillary and ventrolateral (vs. absence). Leptobrachella murphyi sp. nov. differ from L. minima by having a relatively smaller body size in males (SVL 23.2–24.9 mm vs. 25.7–31.4 mm), wide lateral fringes on toes (vs. absent), dorsal skin shagreened with reddish tubercles and folds (vs. mostly smooth) and iris that is orange above and silver white below (vs. dark gold above and grey below). The new species differs from L. zhangyapingi by having a smaller body size in males (SVL 23.2–24.9 mm vs. 45.8−52.5 mm) and creamy white belly with small black spots on the margin (vs. throat, chest, and belly nearly immaculate white). In addition, L. murphyi sp. nov. further differs from L. pelodytoides, the notoriously “widespread” species of Leptobrachella in Indochina, by having a relatively smaller body size (SVL 23.2–24.9 mm in males, 29.3–32.1 mm in females vs. 27.5–32.3 mm, 35.5–37.8 mm), small pectoral glands (vs. large), distinct dermal ridges under toes (vs. indistinct), ventral surface creamy white with small black spots on the margin (vs. whitish), wide lateral fringes on toes (vs. narrow). During the final revisions of our paper, two new species of Leptobrachella (L. shiwandashanensis and L. graminicola) were described. Morphologically, L. murphyi sp. nov. can be easily distinguished from L. shiwandashanensis and L. graminicola. Leptobrachella murphyi sp. nov. differs from L. shiwandashanensis by having a relatively smaller body size (SVL 23.2–24.9 mm in males, 29.3–32.1 mm in females vs. 26.8–29.7 mm, 33.7–35.9 mm), toe webbing rudimentary and lateral fringes wide (vs without webbing and lateral fringes on toes), heels meeting when the tibias perpendicular to the body axis (vs. not meeting). Leptobrachella murphyi sp. nov. differs from L. graminicola by having dermal ridges on the dorsal surface (vs. dorsal surface lacking dermal ridges), as well as the absence of a dark brown throat with light grey-brown flecks and spots (vs. presence), the absence of a row of large white spots on the outer edge of the tarsus extending from the heel to the inner metatarsal tubercle (vs. presence). Advertisement call comparisons. The advertisement call of L. murphyi sp. nov. (Results and Fig. 3) differs from all other congeners occurring north of the Isthmus of Kra for which comparable acoustic data are available in consisting of uniform and continuous calls with four to five pulses in each note on average and a peak frequency of 4.5–4.7 kHz. Of the congeners in the region with known calls, the new species can be separated from L. purpurus, L. tuberosus, L. puhoatensis and L. yingjiangensis by not having an invari, Published as part of Chen, Jin-Min, Suwannapoom, Chatmongkon, Wu, Yun-He, Poyarkov, Nikolay A., Xu, Kai, Pawangkhanant, Parinya & Che, Jing, 2021, Integrative taxonomy reveals a new species of Leptobrachella (Anura: Megophryidae) from the mountains of northern Thailand, pp. 191-214 in Zootaxa 5052 (2) on pages 200-210, DOI: 10.11646/zootaxa.5052.2.2, http://zenodo.org/record/5568539, {"references":["Ohler, A., Wollenberg, K. C., 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","Chen, J. M., Poyarkov, Jr. N. J., Suwannapoom, C., Lathrop, A., Wu, Y. H., Zhou, W. W., Yuan, Z. Y., Jin, J. Q., Chen, H. M., Liu, H. Q., Nguyen, T. Q., Nguyen, S. N., Van, D. T., Eto, K., Nishikawa, K., Matsui, M., Orlov, N. L., Stuart, B. L., Brown, R. M., Rowley, J. J. L., Murphy, R. W., Wang, Y. Y. & 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","Taylor, E. H. (1962) The amphibian fauna of Thailand. The University of Kansas science bulletin, 63, 265 - 599. https: // doi. org / 10.5962 / bhl. part. 13347","Yuan, Z. Y., Sun, R. D., Chen, J. M., Rowley, J. J. L., Wu, Z. J., Hou, S. B., Wang, S. N. & 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","Yang, J. H., Zeng, Z. C. & Wang, Y. Y. (2018) Description of two new sympatric species of the genus Leptolalax (Anura: Megophryidae) from western Yunnan of China. PeerJ, 6, 1 - 32. https: // doi. org / 10.7717 / peerj. 4586"]}

The genus Leptobrachella is one of the most speciose and taxonomically troubling groups of Asian anurans. Herein, we describe a new species of Leptobrachella from Doi Inthanon, Chiang Mai Province, Thailand based on the integration of morphological, acoustic and molecular data. The new species, Leptobrachella murphyi sp. nov., is morphologically distinguished from its congeners on the basis of body size, dorsal skin texture and coloration, ventral coloration and pattern, degree of webbing and fringing on the fingers and toes, iris coloration in life, pattern of markings on flanks and pattern of body macroglands. It also differs from its phylogenetically close congeners by an uncorrected p-distance of >9.6% for a fragment of 16S rRNA. The advertisement call of the new species consists of 4.5–4.7 kHz (at 15 °C) and without a distinct introductory note. Leptobrachella murphyi sp. nov. likely occurs across the Thanon Thong Chai Range and analyses provide evidence of unknown biodiversity and species composition on Doi (mountain) Inthanon. In addition, the congeneric species L. minima was also confirmed in Doi Inthanon. The coexistence pattern of Leptobrachella in Doi Inthanon deserves further study. As Thailand’s highest mountain and biodiversity reservoir, the need for further biological exploration is urgent given ongoing habitat loss and degradation.

We described a new species of the genus Leptobrachella from Southern Guangxi, China, based on morphological characteristics, molecular analyses and bioacoustics. This new species, Leptobrachella shiwandashanensis sp. nov., occurs in sympatry with Leptobrachella shangsiensis. However, phylogenetic analyses based on the mitochondrial 16S gene fragment revealed that the new species was clustered in a monophyletic group and a considerable genetic divergence existed between the new species and L. shangsiensis (p-distance: 7.9%) as well as its congeners (minimum p-distance: 7.0%). The new species also differed from its congeners by a combination of the following characteristics: (1) small size (SVL 26.8–29.7 mm in males; 33.7–35.9 mm in females); (2) pale brown dorsal surfaces, with a brown inverse-triangle-shaped marking between the eyes; (3) creamy white ventral surface with brown spots on the lateral margin, with a near immaculate creamy white throat and chest; (4) without webbing and lateral fringes on toes; (5) flanks with irregular black spots; (6) tibia-tarsal articulation reaching the posterior of the eye in males but reaching the shoulder in females; (7) heels that do not meet when the thighs are appressed at right angles to the body; (8) bicolored iris, with the upper half brownish-red, fading to silver in the lower half; and (9) a call consisting of a single note and a dominant frequency of 5.3–5.7 kHz (recorded at 23°C).

The complete mitochondrial genome of the Leptobrachella alpina Fei, Ye, and Li 1990, was assembled for the first time. The mitogenome of this species was 17,763 bp in length, containing 13 protein-coding genes, two ribosomal RNA genes (12S rRNA and 16S rRNA), 22 transfer RNA genes (tRNA), and a non-coding control region (D-loop). The base content of the mitogenome was that A, T, G, and C occupied 28.5%, 30.8%, 15.1%, and 25.6%, respectively. The phylogenetic analysis was conducted based on 17 complete mitogenome sequences of the family Megophryidae by the Bayesian inference approach. The phylogenetic tree suggested that Leptobrachium and Oreolalax clustered into a clade and formed a sister group with Leptobrachella. This work is critical for the further genetic research and conservation of this species.

We describe the tadpole of the Botsford’s leaf-litter frog Leptobrachella botsfordi, a species endemic to Mount Fansipan in northwest Vietnam, for the first time. Tadpoles of this species were found in steep, fast-flowing streams at elevations between 2500–2600 m asl. We also report a previously unknown locality for this species on Mount Fansipan, which increases the species’ known Extent of Occurrence from 8 km2 to 36 km2. Our findings will inform subsequent conservation initiatives for this poorly known and highly threatened species.

The species-rich, megophryid frog genus Leptobrachella Smith, 1925 is distributed throughout the uplands of mainland Southeast Asia but is conspicuously absent from the Cardamom Mountains of southwestern Cambodia, where it has been known only by a single, undetermined metamorphic specimen collected nearly two decades ago. We report two adult female specimens of Leptobrachella collected a decade later at a second locality in the Cardamom Mountains and use mitochondrial DNA, nuclear DNA, and morphology to show that it represents a distinct species. Leptobrachella neangi sp. nov., named after its collector, is most closely related in mitochondrial and nuclear DNA to L. fuliginosa (Matsui, 2006) and L. melanoleuca (Matsui, 2006) from western Thailand, but has uncorrected pairwise distances of 8.69–10.99% in a mitochondrial 16S gene fragment from its two sister species. The new species is also readily distinguished from these and other congeners by having the combination of (1) SVL 35.4–36.3 mm in two adult females, (2) distinct dorsolateral glandular line absent, (2) belly transparent, immaculate purplish gray in life, creamy white in preservative, (3) dark inguinal blotch absent, (4) tympanum with black coloration extending from line under supratympanic fold, (5) dorsal skin with small, irregular bumps and ridges, and (6) iris coppery orange around pupil, fading to gold at periphery, not distinctly bicolored. The new species is the first named Leptobrachella from the Cardamom Mountains and the third from Cambodia.

A new species of Leptobrachella, Leptobrachella wulingensis sp. nov. is described based on specimens collected from the Wuling mountains in Hunan Province, China. The new species can be distinguished from its congeners by a combination of having a SVL body size range of 24.5–32.8 mm in four adult males and 29.9–38.5 mm in three adult females; dorsal surface brown to reddish brown with indistinct markings; ventral surface creamy white, often with pale brown speckling on chest and margins; flanks with small to moderate black spots; skin on dorsum shagreened with sparse large warts, sometimes with short longitudinal ridges; toes with rudimentary webbing and narrow lateral fringes; dermal ridges under toes interrupted at the articulations; and iris bicolored with a bright orange or golden upper half, fading to silver in the lower half. Uncorrected sequence divergence between L. wulingensis sp. nov. and homologous 16S rRNA sequences available for all known species in the genus are ≥ 2.3%–2.9%.

Leptobrachella wulingensis sp. nov. Figs. 3���6. Holotype. CSUFT 201, adult female, collected by Tianyu Qian and Xin Xia from Tianzishan Nature Reserve, Zhangjiajie, Hunan Province, China (29��23���12.67���N, 110��27���29.6���E), elevation 979 m. on September 9, 2019. Paratypes. CSUFT 177, adult female, CSUFT 178, subadult female and CSUFT 179, 180, two adult males, collected by Tianyu Qian and Xin Xia from Tianquanshan Forest Park, Zhangjiajie, Hunan Province, China (29��15���57.58���N, 110��12���13.38���E), elevation 414 m. on September 7, 2019; CSUFT 193, adult male and CSUFT 194, adult female, collected by Tianyu Qian and Xin Xia from Tianquanshan Forest Park (29��15���39.43���N, 110��13���29.47���E), elevation 530 m. on September 8, 2019; CSUFT 200, adult male, CSUFT 202���204, three subadult females, collected by Tianyu Qian and Xin Xia from the same location as holotype, on September 9, 2019; CSUFT 301, subadult female and CSUFT 302, 303, two subadult males, collected by Yue Cao and Xin Xia from the same location as the holotype, on August 6, 2019. Etymology. The specific name wulingensis is derived from the Wuling Mountains, where the two collection localities occur. The suggested common name is the ���Wuling leaf-litter toad��� in English and ���W�� L��ng Zh��ng T&umacr; Ch��n (&Emacr;������������)��� in Chinese. Diagnosis. Leptobrachella wulingensis sp. nov. can be distinguished from its congeners by having (1) medium body size (SVL 24.5���32.8 mm in four adult males, 29.9���38.5 mm in three adult females); (2) dorsum brown to reddish brown with indistinct markings; (3) dorsal skin shagreened with sparse, large warts, sometimes with longitudinal ridges; (4) ventral surface of body creamy white or translucent creamy white, with distinct or indistinct pale brown speckling on chest and margins, and marbled texture on belly; small tubercle-like white dots presents on chest and anterior region of abdomen in some specimens; (5) tympanum distinct, slightly concave; (6) flanks with small to moderate black spots, with glandular warts present among these spots; (7) absence of webs and lateral fringes on fingers, toes with rudimentary webbing and narrow lateral fringes; (8) distinct longitudinal ridges under toes and interrupted at the articulations; (9) tibia length 44%���49% of SVL in males; (10) iris bicolored with bright orange or golden upper half, fades to silver in lower half; (11) ventrolateral glands distinctly visible, forming an incomplete line; small supra-axillary and femoral glands present; pectoral glands indistinct; (12) tongue cordiform, notched behind; (13) and dense small white conical spines present on lateral and ventral surface of tarsus, surface of tibia-tarsal, inner-side surface of shank and around cloacal region. Description of holotype. Adult female, medium in body size, SVL 31.1 mm; head length about equal to head width, HDL/HDW 1.03; snout rounded, snout length equal to eye diameter, SNT/EYE 1.00; nostril closer to snout than eye; canthus rostralis gently rounded; loreal region slightly concave; pupil vertical; vomerine teeth absent; tympanum rounded, slightly concave, upper margin in contact with supratympanic ridge; supratympanic fold from eye towards supra-axillary gland; tongue cordiform, notched behind. Tips of fingers rounded, slightly swollen; relative finger lengths III> IV = II> I; subarticular tubercles absent; inner palmar tubercle large and rounded, connected with a smaller, round outer palmar tubercle; absence of webbing and lateral fringes on fingers; tips of toes rounded and thickened; relative toe length IV> III> V> II> I; subarticular tubercles absent, replaced by longitudinal dermal ridges, extending on phalanges and interrupted at the articulations; elongate, large oval inner metatarsal tubercle present; toes webbing rudimentary; narrow lateral fringes present on all toes. Tibia 47% of snout-vent length; tibiotarsal articulation reaches to middle of eye. Dorsal surface shagreened, the region between eyes relatively smooth, dense large warts present on the posterior part of dorsum; upper eyelids and limbs with small tubercles; flanks with distinct glandular warts forming two rows; skin on ventral surfaces of trunk, head and forelimbs smooth; white conical spines present on lateral and ventral surface of tarsus, surface of tibia-tarsal, inner-side surface of shank and surface around cloacal region; pectoral gland elongate, and femoral gland oval; pectoral glands greater than femoral glands, PEC/FEM 1.08; femoral glands situated on the posteroventral surface of thigh, closer to knee than to vent; supra-axillary gland raised; ventrolateral gland line distinctly visible. Coloration of holotype in life. Dorsal surface brown with a dark inverted triangular marking in the interorbital region and followed by a "W" shaped mark between axillae; a ���Y��� shape mark between the loreal region linked to the triangular marking between the eyes; three vertical bars present at the rostral-eye region; supratympanic line weak, lower edge black; granules present on dorsum, flanks and limbs having dark orange pigmentation in the center; moderate black spots presents on flanks; glandular warts on flanks white to orange; transverse bars presents on lower arms and legs, as well as fingers and toes; elbow and upper arms coppery orange. Ventral surface of throat and limbs greyish-pink, chest greyish pink with creamy white pigmentation, belly creamy white but translucent with greyish-pink skin color; dense small tubercle-like white dots present on chest; marbled texture present on belly; vaguely visible brown flecks present on chest and margin of belly, close to the ventrolateral gland line; supra-axillary gland orange, ventrolateral glands whitish orange, femoral and pectoral glands white; iris bicolored, golden in the upper half, and fade to dark silver in lower half. Coloration of holotype in preservative. The background color on the dorsum faded to dark grey; dorsalateral markings have become hardly visible; dark vertical bars, transverse bars and black spots distinct; the orange color on tubercles, glands, and elbow has faded to white-grey; ventral surfaces of body and limbs grey-white with dark pigmentations present at the edge of jaw and margin of belly, sparse pigmentations vaguely visible on the chin and chest; iris uniformly dark grey, the upper half and lower half are indistinguishable. Measurements of holotype (in mm). SVL 31.1; HDL 10.8; HDW 10.5; SNT 4.2; EYE 4.2; IOD 3.4; TMP 1.7; TEY 1.1; TIB 14.5; ML 7.9; PL 13.1; PEC 1.4; FEM 1.3. Comparison. Comparative morphological data of Leptobrachella wulingensis sp. nov., and 53 recognized Leptobrachella species occurring north of the Isthmus of Kra are listed in Table 3. Genetically, L. wulingensis sp. nov. is most closely related to L. alpina, L. bijie, L. bourreti, L. eos, L. oshanensis, L. purpuraventra, L. purpurus, and L. suiyangensis. The uncorrected 16S rRNA between L. wulingensis sp. nov. and above mentioned species are 3.6%���4.6% (L. alpina), 2.3%���2.9% (L. bourreti), 4.0% (L. eos), 3.9%���4.3% (L. oshanensis), 4.0% (L. purpuraventra), 4.3% (L. purpurus), and 4.3% (L. suiyangensis). Morphological characters can further distinguish them by the following: L. wulingensis sp. nov. differs from the phylogenetically close congener, L. purpurus by having narrow lateral fringes on toes (vs. wide in purpurus), brown/reddish brown above in life (vs. purplish brown in purpurus), presence of longitudinal skin folds on dorsum (vs. absent in purpurus), supratympanic line weak (vs. distinct and black in purpurus), and dorsal tubercles orange in life (vs. reddish in purpurus). L. wulingensis sp. nov. differs from the phylogenetically close congener, L. alpina by dorsum without white tiny flecks (vs. distinct white tiny flecks present on dorsum in alpina, see Yang et al., 2018: Fig. 6C), absence of distinct dark brown spots/blotches on belly (present in alpina, see Yang et al., 2018: Fig. 6D) in life, tibiotarsal articulation reaches to middle of the eye (vs. reaches to anterior corner of the eye in alpina), narrow lateral fringes on toes (vs. wide in males in alpina), and indistinct dorsolateral markings (vs. distinct in alpina). L. wulingensis sp. nov. differs from the phylogenetically close congener, L. bourreti by having a relatively smaller body size (males SVL 24.5���32.8 mm vs. 28.0��� 36.2 mm in bourreti; females SVL 29.9���38.5 mm vs. 41.8��� 45.0 mm in bourreti), dermal ridges under toes distinct (vs. indistinct in bourreti), absence of dermal fringes on fingers (vs. present on Fingers II and III in bourreti), head longer than or as long as wide (vs. head wider than long in bourreti), indistinct dorsolateral markings (vs. distinct in bourreti), and indistinct supra-axillary glands in specimens reaching metamorphosis (see Fig. 6A) (vs. distinct in bourreti). L. wulingensis sp. nov. differs from the phylogenetically close congener, L. eos by having narrow lateral fringes on toes (vs. wide in eos), presence of black spots on flanks (vs. absent in eos), large warts present on dorsal surface (vs. absent in eos), orange dorsal surface of elbow and upper arms in life (vs. reddish-brown in eos), and ventral surface creamy white with distinct/indistinct brown speckling in life (vs. throat with orange/yellow spots, chest and belly pearly-orange in eos). L. wulingensis sp. nov. differs from the phylogenetically close congener, L. oshanensis by the presence of webbing and lateral fringes on toes (vs. absent in oshanensis), longitudinal ridges under toes interrupted at the articulations (vs. not interrupted in oshanensis), large warts present on dorsal surface (vs. absent in oshanensis), supratympanic line weak (vs. distinct and black in oshanensis), pectoral gland lager than femoral gland (vs. reversed condition in oshanensis), femoral gland closer to knee than to vent (vs. reversed condition in oshanensis), and indistinct dorsolateral markings (vs. distinct in oshanensis). L. wulingensis sp. nov. differs from the phylogenetically close congener, L. suiyangensis by having large warts present on dorsal surface (vs. absent in suiyangensis), brown/reddish brown above in life (vs. greyish brown in suiyangensis), supratympanic line weak (vs. deep and black in suiyangensis), indistinct dorsolateral markings (vs. distinct in suiyangensis), upper parts of iris bright orange/golden in life (vs. coppery in suiyangensis), and smaller tympanum (TMP/EYE ratio 0.32���0.52 vs. 0.42���1.06 in suiyangensis). L. wulingensis sp. nov. differs from the phylogenetically close congener, L. purpuraventra and L. bijie by having large warts present on dorsal surface (vs. absent in purpuraventra and bijie), supratympanic line weak (vs. distinct and black in purpuraventra and bijie), ventral surface creamy white with/without brown speckling in life (vs. grey purple with nebulous greyish speckling in purpuraventra and white with nebulous greyish speckling in bijie), longitudinal ridges under toes interrupted at the articulations (vs. not interrupted in purpuraventra and bijie), indistinct dorsolateral markings (vs. distinct in purpuraventra and bijie), and absence of tiny conical spines on surface of chest in males during breeding season (vs. present in purpuraventra and bijie). Compared with the members from the L. applebyi species group. L. wulingensis sp. nov. differs by having a creamy white chest and belly in life (vs. reddish brown or greyish violet in all members, except rowleyae), presence of webbing and lateral fringes on toes (vs. absent in ardens, kalonensis, maculosa, pallida, rowleyae and tadungensis), bicolored iris (vs. uniform in ardens and tadunensis), distinct ventrolateral glandular line (vs. indistinct or absent in macrops, pyrrhops, rowleyae, tadungensis and melica), dorsal skin texture shagreened with sparse large warts (vs. mostly smooth in applebyi, bidoupensis, kalonensis, maculosa and melica), supratympanic line weak (vs. distinct and black in ardens, kalonensis, maculosa and tadungensis), upper parts of iris bright orange/golden in life (vs. reddish or copper in bidoupensis, maculosa and pallida), and lower parts of iris silver in life (vs. gold, golden green, or silver green in macrops, maculosa, pallida, pyrrhops and rowleyae); and further differs from L. applebyi by having a larger body size (males SVL 24.5���32.8 mm vs. 19.6���20.8 mm in applebyi) and the presence of dermal fringes on toes (vs. absent in applebyi); from L. bidoupensis by having a larger body size (males SVL 24.5���32.8 mm vs. 23.6���24.6 mm in bidoupensis) and the presence of glandular warts on flanks (vs. absent in bidoupensis); from L. macrops by the presence of longitudinal ridges on dorsum (vs. absent in macrops) and lateral fringes on toes (vs. absent in macrops); from L. melica by the absence of dermal fringes on fingers (vs. present in melica) and the presence of lateral fringes on toes (vs. absent in melica). Compared with a congener occurring in Hunan province (L. mangshanensis), L. wulingensis sp. nov. differs by having narrow lateral fringes on toes (vs. weak in mangshanensis), distinct longitudinal ridges under toes interrupt- ed at the articulations (vs. longitudinal ridges indistinct and not interrupted at the articulations in mangshanensis), supratympanic line weak (vs. distinct and black in mangshanensis), tibiotarsal articulation reaches to middle of the eye (vs. reaches anterior margin of snout in mangshanensis), dorsal skin texture shagreened with sparse large warts (vs. mostly smooth in mangshanensis), and indistinct dorsolateral markings (vs. distinct in mangshanensis). Compared with three congeners recently described by Chen et al. (2020) from Yunnan Province, L. wulingensis sp. nov. differs from L. niveimontis by having a creamy white chest and belly (vs. marbled with distinct irregular black speckling on bluish-white background in niveimontis), orange supra-axillary glands (vs. white in niveimontis), orange dorsal surface of elbow and upper arms (vs. reddish-brown in niveimontis) in life, and longitudinal ridges under toes interrupted at the articulations (vs. not interrupted in niveimontis); from L. flaviglandulosa by having small white spines around cloacal region (vs. absent in flaviglandulosa), orange supra-axillary glands (vs. yellowish in flaviglandulosa), orange dorsal surface of elbow and upper arms (vs. yellowish in flaviglandulosa), greyish-pink ventral surface of throat and limbs (vs. dark brown in flaviglandulosa), and lacking yellow markings on dorsum in scapular region (vs. present in flaviglandulosa) in life; from L. feii by weak supratympanic line (vs. distinct and black in feii), absence of distinct black blotches scattered on chest and belly (vs. present in feii), longitudinal ridges under toes interrupted at the articulations (vs. not interrupted in feii), small to moderate black spots on flanks (vs. large blotches in feii), and absence of nuptial spines on the dorsal surfaces of first and second fingers (vs. presence in males in feii). For the other congeners known from China, L. wulingensis sp. nov. differs by having narrow lateral fringes on toes (vs. broad or wide in laui, liui, yingjiangensis and yunkaiensis; absent in nyx and ventripunctata), absence of lateral fringes on fingers (vs. present in laui and yingjiangensis), supratympanic line weak (vs. distinct and black in maoershanensis, shangsiensis, ventripunctata and wuhuangmontis), longitudinal ridges under toes interrupted at the articulations (vs. not interrupted in laui, maoershanensis, wuhuangmontis and yunkaiensis), indistinct dorsalateral markings (vs. distinct in liui, maoershanensis, pelodytoides, tengchongensis, ventripunctata, wuhuangmontis and yunkaiensis), absence of black spots or dark blotches on ventral surface (vs. present in maoershanensis, tengchongensis and ventripunctata), black spots on flanks small to moderate in size (vs. large in maoershanensis, pelodytoides and tengchongensis), distinct dermal ridges under toes (vs. indistinct in nyx and pelodytoides) and distinct ventrolateral glandular line (vs. indistinct in nyx and tengchongensis); and further differs from L. laui by the presence of longitudinal ridges on dorsum (vs. absent in laui); from L. liui by the absence of light dorsal markings in preservatives (vs. large, dark dorsal markings edged with light margins in liui, see Sung et al., 2014: Fig. 4B); from L. shangsiensis by having shagreened dorsal skin texture with sparse large warts (vs. smooth with numerous tiny tubercles in shangsiensis); from L. pelodytoides by having distinct dermal fringes under toes (vs. indistinct in pelodytoides); from L. tengchongensis by having bicolored iris (vs. uniform in tengchongensis); from L. ventripunctata by upper parts of iris bright orange/golden in life (vs. copper in ventripunctata) and lower parts of iris silver in life (vs. grey-brown in ventripunctata); from L. yingjiangensis by having orange supra-axillary glands in life (vs. brown in yingjiangensis) and the absence of white and dark brown mottling on ventral surface of thigh (vs. present in yingjiangensis); from L. yunkaiensis by having creamy white ventral surface (vs. pinkish in yunkaiensis) in life. For the other 18 congeners occurring north of the Isthmus of Kra, L. wulingensis sp. nov. differs by having indistinct dorsalateral markings (vs. distinct in botsfordi, fuliginosa, khasiorum, lateralis, minima, nahangensis, nokrekensis, pluvialis, puhoatensis, sungi, tamdil and zhangyapingi), presence of black spots on flanks (vs. absent in aerea, botsfordi, firthi and tuberosa), creamy white ventral surface (vs. reddish brown or bright orange in botsfordi, puhoatensis and, Published as part of Qian, Tian-Yu, Xia, Xin, Cao, Yue, Xiao, Neng-Wen & Yang, Dao-De, 2020, A new species of Leptobrachella (Anura: Megophryidae) Smith, 1925 from Wuling Mountains in Hunan Province, China, pp. 491-526 in Zootaxa 4816 (4) on pages 497-505, DOI: 10.11646/zootaxa.4816.4.4, http://zenodo.org/record/3954622, {"references":["Yang, J. H., Zeng, Z. C. & Wang, Y. Y. (2018) Description of two new sympatric species of the genus Leptolalax (Anura: Megophryidae) from western Yunnan of China. PeerJ, 6, e 4586. https: // doi. org / 10.7717 / peerj. 4586","Chen, J. M., Xu, K., Poyarkov, N. A., Wang, K., Yuan, Z. Y., 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 (3), 292 - 313. https: // doi. org / 10.24272 / j. issn. 2095 - 8137.2020.036","Sung, Y. H., Yang, J. H. & Wang, Y. Y. (2014) A new species of Leptolalax (Anura: Megophryidae) from southern China. Asian Herpetological Research, 5 (2), 80 - 90. https: // doi. org / 10.3724 / SP. J. 1245.2014.00080","Rowley, J. J. L., Hoang, D. H., Le, T. T. D., Dau, V. Q. & Cao, T. T. (2010 c) A new species of Leptolalax (Anura: Megophryidae) from Vietnam and further information on Leptolalax tuberosus. Zootaxa, 2660 (1), 33 - 45. https: // doi. org / 10.11646 / zootaxa. 2660.1.3","Mo, X., Shen, Y., L, Y. & Wu, X. (2010) A new species of Megophrys (Amphibia: Anura: Megophryidae) from the northwestern Hunan Province, China. Current Zoology, 56 (4), 432 - 436. https: // doi. org / 10.1093 / czoolo / 56.4.432"]}

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

This study describes a new species of the genusLeptobrachella,Leptobrachella suiyangensissp. nov.from the Huoqiuba Nature Reserve, Suiyang County, Guizhou Province, China, based on morphological data and phylogenetic analyses (16S rRNA mtDNA). The new species can be distinguished from other congeners by the molecular divergence and by a combination of morphological characters, including body size, dorsal and ventral patterns, dorsal skin texture, size of the pectoral and femoral glands, degree of webbing and fringing on the toes and fingers, dorsum coloration, and iris coloration in life. Currently, the genusLeptobrachellacontains 75 species, 21 of which are found in China, including seven species reported from Guizhou Province. The uncorrected sequence divergence percentage betweenLeptobrachella suiyangensissp. nov.and all homologous DNA sequences available for the 16S rRNA gene was found to be >4.7%. The new record of the species and its relationships with others in the same genus imply that species distribution, habitat variation, environmental adaptation, and diversity of the genusLeptobrachellain southwest China need to be further investigated.

Two new species of the dwarf litter frog genus Leptobrachella are described from North and Central Kalimantan, Indonesian part of the Borneo Island. Leptobrachella fusca n. sp. from Bulungan Regency, differs from all congeneric species in the following combination of characters: body small (SVL: 16.3 mm in a male); dorsum uniformly dark brown; sides of body without marking; a series of elongate ventrolateral glands present on flank; ventrum darkly pigmented except for throat and posterior portion; lineae masculinae invisible through ventral skin in preservative; and toe webs rudimentary. Leptobrachella bondangensis n. sp. from Gunung (=Mt.) Bondang of Murung Raya Regency, is distinct in possessing the following combination of characters: body large as this genus (SVL: 17.8 mm in a male); dorsum grayish brown; sides of body without dark blotches; a sparse series of rounded ventrolateral glands present on flunk; ventrum pale gray with dark mottles; lineae masculinae invisible through ventral skin in preservative; and toe webs rudimentary.

We studied specimens of the genus Leptobrachella Smith, 1925 collected from West Kalimantan, Indonesian part of the island of Borneo, and housed in the Bogor Museum as L. mjobergi Smith, 1925. Our morphological examination proved that the specimens should be identified as L. juliandringi Eto, Matsui et Nishikawa 2015. This is the first record of L. juliandringi from outside Sarawak, Malaysian Borneo. Brief discussion is made on the distribution of the genus in Borneo.

The family Megophryidae is one of the most poorly known groups of amphibians in Son La Province, Vietnam. In the herpetofaunal list of Vietnam in 2009 only two species of megophryids were recorded from this province and recent studies have documented five additional species of Megophryidae from Son La Province. Based on recent field work in northern Vietnam, we report six new provincial records of the family Megophryidae from Son La Province, namely Leptobrachium ailaonicum, Leptobrachella sungi, Megophrys feae, M. jingdongensis, M. microstoma and M. parva. In addition, morphological data were provided for each species, based on newly collected specimens. Our findings bring the species number of the family Megophryidae to 13 in Son La Province.

The advertisement calls of five males were recorded in the field at Suichang, Zhejiang, China (28��22���14��� N, 118��53���25��� E) in April 2017. Calls were recorded using a Sony IC ICD-SX1000 recorder with an internal microphone placed at a distance of approximately 0.5 m from focal males, with a sampling frequency resolution of 44.1 kHz and 16- bit resolution. A total of 245 calls were recorded and the number of calls ranged from 22 to 81 among individuals. The air temperature at the time of recording varied between 14.7���20.2�� C. Calls were analyzed using Praat 6.0.36. Stream background noise was reduced in Cool Edit Pro 2.1. Description and terminology of call parameters followed (Xu et al. 2005): call duration (CD, in s), note number in a single call (NN), call intensity (CTI, in dB), dominant frequency (DF, in Hz), fundamental frequency (F0, in Hz), the frequency of first to third formants (F1, F2 and F3, in Hz). We used withinmale coefficients of variation (CV W) of each parameter in order to classify them as dynamic (CV W > 10%), static (CV W 1.0 are usually indicative that the parameter could be used for individual recognition (e.g., Gambale et al. 2014). Values are presented as mean �� standard deviation and range. The advertisement call of L. liui (Fig. 1) consists of multiple pulsed notes, and is highly amplitude-modulated, with amplitude peaking towards the start of each call. Calls of the five males consisted of 3.11 �� 0.35 notes (range 2���4 notes), with one pulse per note, CD of 0.063 �� 0.008 s (range 0.026 ��� 0.078 s), CTI of 72.04 ��1.96 dB (range 67.63���79.09 dB). The DF ranged from 4814.5 to 5489.6 Hz (average 5234.4 �� 143.4 Hz), whereas the F0 ranged from 811.8 to 1179.5 Hz (average 968.0 �� 73.7 Hz). The F1, F2 and F3 were emitted from 1693.4 to 2419.9 Hz (average 2050.9 �� 150.5 Hz), 2632.0 to 4049.7 Hz (average 3215.8 �� 391.4 Hz) and 4161.9 to 5258.3 Hz (average 4635.3 �� 267.6 Hz), respectively. The CV W of CD, NN, CTI, DF, F0, F1, F2 and F3 were 9.6 �� 4.1 % (range 4.0���15.5 %), 6.2 �� 6.7 % (range 0���16.1 %), 1.9 �� 0.5 % (range 1.2���2.3 %), 1.8 �� 0.9 % (range 0.9���3.2 %), 7.1 �� 1.9 % (range 4.3���9.7 %), 5.7 �� 2.3 % (range 3.1���9.3 %), 6.3 �� 3.5 % (range 3.0���9.3 %) and 2.7 �� 1.0 % (range 1.4���3.7 %), respectively. The ratios of CV A /CV W in CD, NN, CTI, DF, F0, F1, F2 and F3 were 1.3, 1.8, 1.5, 1.4, 1.1, 1.2, 1.9 and 2.2, respectively. The advertisement call of L. alpinus contains approximately nine notes, with average CD of 0.270 �� 0.069 s, average DF, F0 and F1 all of 6700 Hz (Xu et al. 2005). The advertisement call of L. oshanensis consists of three notes, with average CD of 0.046 �� 0.013 s (range 0.029 ���0.067 s) and DF ranging between 4402���4633 Hz (average 4521.9 �� 79.6 Hz) (Jiang et al. 2002). These reported call parameters are different among the three Chinese endemic Leptobrachella species, and could be helpful in species identification in the field. CTI, DF and F3 are classified as static traits in L. lilui, whereas other parameters are intermediate due to their average CV W between 5 % and 10 %. The ratios of CV A /CV W in all parameters were more than 1.0, suggesting that advertisement calls of L. liui could play a role in individual recognition., Published as part of Ding, Guo-Hua, Chen, Zhi-Qiang, Tang, Yun & Lin, Zhi-Hua, 2019, The advertisement call of Leptobrachella liui Fei and Ye, 1990 (Anura: Megophryidae), pp. 588-590 in Zootaxa 4576 (3) on pages 588-589, DOI: 10.11646/zootaxa.4576.3.11, http://zenodo.org/record/3993508, {"references":["Xu, J. X., Xie, F., Jiang, J. P., Mo, Y. M. & Zheng, Z. H. (2005) The acoustic features of the mating call of 12 anuran species. Chinese Journal of Zoology, 40, 12 - 19.","Gambale, P. G., Signorelli, L. & Bastos, R. P. (2014) Individual variation in the advertisement calls of a Neotropical treefrog (Scinax constrictus). Amphibia-Reptilia, 35, 271 - 281. https: // doi. org / 10.1163 / 15685381 - 00002949","Jiang, J. P., Xie, F., Fei, L., Ye, C. Y. & Zheng, M. Q. (2002) Mating calls of six forms of pelobatid in Wawu Mountain National Forest Park, Sichuan, China (Anura: Pelobatidae). Zoological Research, 23, 89 - 94."]}

Leptobrachella (Anura: Megophryidae) is currently composed of seventy-one species distributed in China and Southeast Asia (Frost 2018). Fourteen species of Leptobrachella are currently known from southern China: L. alpina (Fei, Ye & Li, 1990), L. laui (Sung, Yang & Wang, 2014), L. liui (Fei & Ye, 1990), L. mangshanensis (Hou, Zhang, Hu, Li, Shi, Chen, Mo & Wang, 2018), L. maoershanensis (Yuan, Sun, Chen, Rowley & Che, 2017), L. oshanensis (Liu, 1950), L. purpura (Yang, Zeng & Wang, 2018), L. tengchongensis (Yang, Wang, Chen & Rao, 2016), L. wuhuangmontis (Wang, Yang & Wang, 2018), L. yingjiangensis (Yang, Zeng & Wang, 2018), L. yunkaiensis (Wang, Li, Lyu & Wang, 2018), L. ventripunctata (Fei, Ye & Li, 1990), L. pelodytoides (Boulenger, 1893) and L. sungi (Lathrop, Murphy, Orlov & Ho, 1998) and the first eleven species are endemic of this region (AmphibiaChina 2018). Many new species in this genus were discovered based on molecular, morphometric and bioacoustic data in recent years (Frost 2018). In comparison to molecular and morphological data, bioacoustics is especially useful in species identification of Leptobrachella in field surveys (Rowley et al. 2016). Although the advertisement calls of many Leptobrachella species were described along with the species descriptions (e.g., Rowley et al. 2016), the advertisement calls of only three Chinese endemic species (L. alpinus; L. laui and L. oshanensis) are known (Jiang et al. 2002; Matsui 2006; Xu et al. 2005). Here, we describe the advertisement calls of L. liui, which inhabits rocky streams in southeastern China (Fujian, Jiangxi and Zhejinag) and is morphologically diagnosed by the combination of a set of external color traits (Fig. 1).

Leptobrachella shangsiensis sp. nov. Figs. 2���3 Holotype. NHMG1704002, adult male (Fig. 2 A���F), collected at Shiwandashan National NR, Guangxi Province, China (22.455�� N, 107.048�� E; alt. 500 m). Collected on 26 April 2017 by Weicai Chen, Yunming Mo, Xiaowen Liao and Shichu Zhou. Paratypes. NHMG1401024���031, NHMG1704003���004, ten adult males, NHMG1401032���034, three adult females, collected at same locality as holotype on 3 Jan. 2014 by Weicai Chen and Yunming Mo. NHMG201604001, one adult female, collected at same locality as holotype on 25 April 2016 by Weicai Chen and Yunming Mo. NHMG1704005, one adult female, collected at same locality as holotype on 26 April 2017 by Weicai Chen and Yunming Mo. Etymology. The specific epithet is in reference to the type locality of Shangsi County, Guangxi Province, China. Suggested English name is Shangsi Leaf Litter Toad. Diagnosis. The specimens were assigned to the genus Leptobrachella on the basis of the following characters: small body size, having an elevated inner metacarpal tubercle, having macro-glands on body (including supraaxillary, femoral and ventrolateral glands), lacking vomerine teeth and having small tubercles on eyelids (Dubois 1980; Lathrop et al. 1998; Delorme et al. 2006). Leptobrachella shangsiensis sp. nov. is distinguished from its congeners by a combination of (1) SVL 24.9���29.4 mm in males (n = 11) and 30.8���35.9 mm in females (n = 5), (2) black supratympanic fold with dark reddish rim, (3) ventrolateral glands distinct, forming a broken line, (4) ventral surface yellowish creamy-white with marble texture, (5) iris copper in the upper and silver in the lower fifth and (6) toe webbing basal and lateral fringes distinct, narrow. Description of holotype. Head width greater than head length (HW / HL = 1.15); snout bluntly rounded in profile and dorsal view, projecting slightly over lower jaw; nostril oval-shaped and closer to tip of snout than eye; canthus rostralis distinct; loreal region distinctly sloping, concave; pupil vertical; diameter of eye less than length of snout (ED / SNT = 0.78); tympanum distinct, rounded, diameter 55% that of eye (TD / ED = 0.55); tympanic rim elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings present, located laterally on floor of mouth; tongue large with shadow notch at posterior tip; supratympanic fold distinct, running from corner of eye to axillary gland (Fig. 2 A���D). Forelimbs. Forelimbs thin, slender; tips of fingers rounded, slightly swollen; relative fingers lengths I Hindlimbs. Hindlimbs slender; tips of toes rounded, slightly swollen, similar those of fingers; relative toe lengths I Skin texture and skin glands. Skin on dorsum mostly smooth with numerous tiny tubercles; upper eyelid with small tubercles; ventral skin smooth; pectoral glands visible; femoral glands oval, approximately 1.1 mm in diameter, located on posteroventral surface of thighs, closer to knee than to vent; supra-axillary gland small, oval, approximately 1.1 mm; ventrolateral glands distinct, forming a broken line (Fig. 2 C, D). Color of holotype in life. Dorsum reddish brown with dark brown markings between eyes and scapular region, and scattered some deep reddish tubercles concentrated on upper eyelid and scapular region; black supratympanic fold with dark reddish rim; canthus rostralis with black spots; lateral surfaces of trunk with creamy whitish spots, scattered with about 5 spots; transverse dark-brown bars present on dorsal surface of thighs, tibia, and lower arms; elbows and upper arms orange with light reddish-brown spots on tubercles; ventral surface yellowish creamy-white with marble texture; ventral surfaces of forelimbs and hindlimbs pinkish milk-white with dense whitish speckling; two distinct blackish-grey spots on edge of upper jaw located from snout to area under eye; supra-axillary gland milk-white; femoral and ventrolateral glands white. Iris copper in the upper and silver in the lower fifth (Fig. 2 A���D). Color of holotype in preservative. In preservation, coloration fades to dark grey-brown on dorsum, forelimbs, and hindlimbs. Transverse dark-brown bars are distinct on forelimbs and hindlimbs. Ventral surface is creamywhite. Margin of throat and ventrolateral surfaces of forelimbs and hindlimbs are pale-brown. Macro-glands including supra-axillary glands, femoral and ventrolateral glands are creamy. Measurements of holotype ( mm): SVL 26.8, HL 8.1, HW 9.3, SNT 4.6, ED 3.8, IOD 2.8, TD 2.1, TED 1.3, IN 3.2, EN 1.8, TIB 12.9, FLL 13.4, THL 14.1, PL 12.0, ML 7.0, FEM 1.1. Variation. Measurements of the type series are shown in Table 2. Females (mean SVL 33.3 �� 2.0, n = 5) have larger body size than males (mean SVL 27.4 �� 1.1, n = 11). In life, all specimens have similar coloration (Fig. 3), but in preservation, some specimens have slightly darker brown compared to holotype. Advertisement call. Call descriptions are on the basis of the calls of the holotype, recorded at an ambient temperature of 21.5��C. Calls of L. shangsiensis sp. nov. consisted of a single note, with call duration 64���69 ms (66.03 �� 1.15, n = 30), call repetition rate 3.4 s and intercall interval 184���289 ms (250.53 �� 31.07, n = 30). The dominant frequency was between 5.5���6.5 kHz, but fundamental frequency and harmonics were not clear (Fig. 4). Ecology and distribution. All specimens were found in an evergreen forest in Shiwandashan National NR located at an elevation of 450���550 m (Fig. 5A). Males were calling on rocks within or adjacent to rocky streams at night between 1930���2400 h. Leptobrachella shangsiensis sp. nov. is only known from Shiwandashan National NR. Comparisons. Leptobrachella shangsiensis sp. nov. differs from all other Leptobrachella species based on morphological, molecular and acoustic data. Phylogenetically, L. shangsiensis sp. nov., L. aereus, L. minimus, L. nahangensis, L. nyx, L. pelodytoides, L. pluvialis, L. ventripunctatus and L. wuhuangmontis form a clade. Genetically, among this clade, the smallest genetic distance is 5.9% (L. shangsiensis sp. nov. and L. nyx) and the largest genetic distance is 10.3% (L. shangsiensis sp. nov. and L. ventripunctatus). However, L. shangsiensis sp. nov. can be distinguished from the above-mentioned species by morphological characters. First, L. aereus differs from L. shangsiensis sp. nov. by having a uniformly bronze iris (vs. iris copper in the upper and silver in the lower fifth), the higher frequency calls (6.2���7.9 kHz at 22.4���25.7��C in L. aereus vs. 5.5���6.5 kHz at 21.5��C in the new species) and absent a distinct black supratympanic line and no black spots on flanks (vs. present a black supratympanic line and black spots on flanks). Leptobrachella minimus differs from L. shangsiensis sp. nov. by having no lateral fringes on toes (vs. present narrow lateral fringes on toes) and iris dark gold above and grey below (vs. iris copper in the upper and silver in the lower fifth). Leptobrachella nahangensis differs from L. shangsiensis sp. nov. by having a large body size (male SVL 40.8 mm vs. SVL 24.9���29.4 mm), a uniformly gold iris (vs. iris copper in the upper and silver in the lower fifth) and no lateral fringes on toes (vs. present narrow lateral fringes on toes). Leptobrachella nyx differs from L. shangsiensis sp. nov. by having a relatively large body size (SVL 26.7���32.6 mm in males and 37.0���41.0 mm in females vs. SVL 24.9���29.4 mm in males and 30.8���35.9 mm in females); lateroventral glandular ridge poorly distinct (vs. distinct); dermal ridges under toes rather poorly distinct (vs. distinct); greyish-brown dorsum with dark regularly set rounded spots (vs. reddish brown dorsum); absent dermal fringe along toe V (vs. present); posterior part of thigh with glandular warts (vs. without glandular warts); HW / HL = 0.95 (vs. HW / HL = 1.15); ED / SNT = 1.00 (vs. ED / SNT = 0.78). Leptobrachella pelodytoides differs from L. shangsiensis sp. nov. by having wide webbing on toes (vs. basal toe webbing in L. shangsiensis sp. nov.), poorly distinct dermal ridges under toes (vs. distinct) and present longitudinal skin folds on dorsum (vs. absent). Leptobrachella pluvialis differs from L. shangsiensis sp. nov. by having a relatively small body size (male SVL 21.3���22.3 mm vs. 24.9���29.4 mm), absent dermal fringes on toes (vs. present), dirty white with dark brown marbling on venter (vs. yellowish creamywhite with marble texture) and dark golden iris (vs. iris copper in the upper and silver in the lower fifth). Leptobrachella ventripunctatus differs from L. shangsiensis sp. nov. by having a bicolored iris, copper above and grey-brown below (vs. iris copper in the upper and silver in the lower fifth), absent lateral fringes on toes (vs. present narrow lateral fringes on toes), chest and belly with dark brown spots (vs. yellowish creamy-white ventral surface with marble texture), present longitudinal skin folds on dorsum (vs. absent) and different dominant frequency (6.1���6.4 kHz at 15.0��C vs. 5.5���6.5 kHz at 21.5��C in L. shangsiensis sp. nov.). Leptobrachella wuhuangmontis differs from L. shangsiensis sp. nov. by having a greyish-white ventral surface mixed by tiny white and black dots (vs. yellowish creamy-white ventral surface with marble texture), distinct dark brown blotches on flanks from groin to axilla (vs. about 5 spots on flanks) and rough skin on dorsum body and limbs with skin ridges and dense conical tubercles (vs. nearly smooth dorsal skin with numerous tiny tubercles). Among the genus Leptobrachella, fifteen species (L. baluensis, L. bondangensis, L. brevicrus, L. fusca, L. itiokai, L. juliandringi, L. lateralis, L. mjobergi, L. natunae, L. nokrekensis, L. palmata, L. parva, L. platycephala, L. serasanae and L. tamdil) have no molecular data. Geographically, these species occur in allopatry with L. shangsiensis sp. nov. Leptobrachella baluensis, L. brevicrus, L. itiokai, L. juliandringi, L. mjobergi, L. palmata, L. parva and L. platycephala come from Malaysia; L. bondangensis, L. fusca, L. natunae and L. serasanae from Indonesia; L. lateralis, L. nokrekensis and L. tamdil from northeast India. Morphologically, L. baluensis (male SVL 14.9���15.9 mm), L. bondangensis (male SVL 17.8 mm), L. brevicrus (male SVL 17.1���17.8 mm), L. fusca (male SVL 16.3 mm), L. itiokai (male SVL 15.2���16.7 mm), L. juliandringi (SVL17.0��� 17.2 mm in males and 18.9���19.1 mm in females), L. mjobergi (male SVL 15.7���19.0 mm), L. natunae (male SVL 17.6 mm), L. palmata (male SVL 14.4���16.8 mm), L. parva (SVL15.0��� 16.9 mm in males and 17.8 mm in female) and L. serasanae (female 16.9 mm) have a small body size (vs. SVL 24.9���29.4 mm in males and 30.8���35.9 mm in females in L. shangsiensis sp. nov.). Furthermore, L. baluensis, L. brevicrus, L. bondangensis, L. fusca, L. itiokai, L. juliandringi, L. mjobergi, L. natunae, L. palmata, L. parva and L. serasanae exhibit an arrow-like projection of digital tip (vs. no arrow-like projection of digital tip). Leptobrachella lateralis has no lateral fringes on toes (vs. present narrow lateral fringes on toes). Leptobrachella nokrekensis has reddish orange irises in the upper (vs. irises copper in the upper and silver in the lower fifth). Leptobrachella platycephalus has a large body size (male SVL 35.1 mm vs. SVL 24.9���29.4 mm) and orange red or reddish brown irises (vs. irises copper in the upper and silver in the lower fifth). Leptobrachella tamdil has wide lateral fringes on toes (vs. present narrow lateral fringes on toes) and wide webbing on toes (vs. basal toe webbing). Leptobrachella shangsiensis sp. nov. occurs in sympatry with L. sungi, but L. sungi has a large body size (male SVL 48.3���52.7 mm vs. SVL 24.9���29.4 mm), gold green iris (vs. iris copper in the upper and silver in the lower fifth), absent a distinct black supratympanic line (vs. present) and absent black spots on flanks (vs. present). Leptobrachella laui and liui have a relatively widespread distribution in China. However, L. laui differs from L. shangsiensis sp. nov. by having no black supratympanic fold (vs. present), moderate dermal fringes on fingers (vs. absent), throat transparent pink with brown dusting along anterior margin (vs. throat yellowish creamy-white with marble texture) and head slightly longer than wide (vs. HW / HL = 1.15). Leptobrachella liui differs from L. shangsiensis sp. nov. by having wide lateral fringes on toes (vs. narrow lateral fringes on toes), creamy-white belly with dark brown spots on chest and margins (vs. yellowish creamy-white belly with marble texture), longitudinal ridges under toes interrupted at the articulations (vs. not interrupted) and greyish-brown dorsal coloration in life (vs. brown or reddish-brown dorsal coloration). From the rest 11 known Leptobrachella species from China, L. shangsiensis sp. nov. differs from L. alpinus by having a different dominant frequency (6.7 kHz at 16��C in L. alpinus vs. 5.5���6.5 kHz at 21.5��C in L. shangsiensis sp. nov.) and wide lateral fringes on toes (vs. present narrow lateral fringes on toes in L. shangsiensis sp. nov.). Leptobrachella bourreti exhibits a copper above and golden below iris (vs. iris copper in the upper and silver in the lower fifth), a relatively large body size (male SVL 28.0��� 36.2 mm vs. SVL 24.9���29.4 mm in L. shangsiensis sp. nov.) and poorly distinct dermal ridges under toes (vs. distinct). Leptobrachella eos has a relatively large body size (male SVL 33.1���34.7 mm in L. eos vs. SVL 24.9���29.4 mm in L. shangsiensis sp. nov.), absent black spots on flanks (vs. present), iris orange above and light gold below (vs. iris copper in the upper and silver in the lower fifth in L. shangsiensis sp. nov.), absent a black supratympanic line (vs. present) and wide lateral fringes on toes (vs. present narrow lateral fringes on toes in L. shangsiensis sp. nov.). Leptobrachella mangshanensis has a bicolored iris with bright orange upper and greyish cream below (vs. iris copper in the upper and silver in the lower fifth in L. shangsiensis sp. nov.) and nearly smooth dorsal skin with some small, orange, tubercles and irregular, dark brown stripes (vs. nearly smooth dorsal skin with numerous tiny tubercles in L. shangsiensis sp. nov.). Leptobrachella maoershanensis has a bicolored iris with orange above and silver below (vs. iris copper in the upper and silver in the lower fifth in L. shangsiensis sp. nov.), creamy-white ventral surface with irregular black spots (vs. yellowish creamy-white ventral surface with marble texture) and shagreened dorsal skin small tubercles and longitudinal folds (vs. nearly smooth dorsal skin with numerous tiny tubercles in L. shangsiensis sp. nov.). Leptobrachella oshanensis has a relatively low dominant frequency (4.4���4.6 kHz at 14��C in L. oshanensis vs. 5.5���6.5 kHz at 21.5��C in L. shangsiensis sp. nov.), absent lateral fringes on toes (vs. present narrow lateral fringes on toes in L. shangsiensis sp. nov.) and present wide webbing on toes (vs. basal toe webbing in L. shangsiensis sp. nov.). Leptobrachella purpurus has a relatively low dominant frequency (4.3��� 4.5 kHz at 15��C in L. purpurus vs. 5.5���6.5 kHz at 21.5��C in L. shangsiensis sp. nov.), a dull white venter with indistinct grey dusting (vs. yellowish creamy-white ventral surface with marble texture in L. shangsiensis sp. nov.) and a bicolored iris with orange yellow upper half and sliver white lower half (vs. iris copper in the upper and silver in the lower fifth in L. shangsiensis sp. nov.). Leptobrachella tengchongensis has a uniformly dark brown iris (vs. iris copper in the upper and silver in the lower fifth in L. shangsiensis sp. nov.), shagreened dorsal skin with small tubercles (vs. nearly smooth dorsal skin with numerous tiny tubercles in L. shangsiensis sp. nov.) and white venter with dark brown blotches (vs. yellowish creamy-white ventral surface with marble texture in L. shangsiensis sp. nov.). Leptobrachella yingjiangensis has a bicolored iris with orange yellow above and sliver white below (vs. iris copper in the upper and silver in the lower fifth in L. shangsiensis sp. nov.), wide lateral fringes on toes (vs. narrow lateral fringes on toes in L. shangsiensis sp. nov.), shagreened dorsal skin with fine, round brown tubercles (vs. nearly smooth dorsal skin with numerous tiny tubercles in L. shangsiensis sp. nov.), ventral surface of body creamy white and scattered with distinct small dark brown flecks on chest and lateral sides of belly (vs. yellowish creamywhite ventral surface with marble texture in L. shangsiensis sp. nov.) and a different dominant frequency (5.7���5.9 kHz at 19��C in L. yingjiangensis vs. 5.5���6.5 kHz at 21.5��C in L. shangsiensis sp. nov.). Leptobrachella yunkaiensis has a pinkish surface of belly with distinct or indistinct light dark brown speckling and black tympanum (vs. yellowish creamy-white belly and brown tympanum in L. shangsiensis sp. nov.) and shagreened dorsal skin with skin ridges and raised warts (vs. nearly smooth dorsal skin with numerous tiny tubercles in L. shangsiensis sp. nov.) Besides the above-mentioned species, the remaining ones (35 of 71 species) belong to different phylogenetic clades, exhibit high genetic variation (ranging from 11.9% to 19.0%) and allopatric distribution. Leptobrachella arayai, L. dringi, L. fritinniens, L. gracilis, L. hamidi, L. heteropus, L. kajangensis, L. kecil, L. marmoratus, L. maurus, L. melanoleucus, L. pictus, L. platycephalus, L. sabahmontanus and L. solus lack supra-axillary and ventrolateral glands (vs. present in L. shangsiensis sp. nov.). The following species, L. applebyi (reddish brown with white specking), L. ardens (reddish brown with, Published as part of Chen, Weicai, Liao, Xiaowen, Zhou, Shichu & Mo, Yunming, 2019, A new species of Leptobrachella (Anura: Megophryidae) from southern Guangxi, China, pp. 67-82 in Zootaxa 4563 (1) on pages 72-79, DOI: 10.11646/zootaxa.4563.1.3, http://zenodo.org/record/2600975, {"references":["Lathrop, A., Murphy, R. W., Orlov, N. & Ho, C. T. (1998) Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Amphibia-Reptilia, 19, 253 - 267. https: // doi. org / 10.1163 / 156853898 X 00160"]}

This study describes a new member of the genus Leptobrachella, Leptobrachella shangsiensis sp. nov., based on morphological, acoustic and molecular analyses from southern Guangxi, China. L. shangsiensis sp. nov. differs from its congeners by a combination of the following characters: (1) SVL 24.9–29.4 mm in 11 males and 30.8–35.9 mm in 5 females, (2) black supratympanic fold with dark reddish rim, (3) ventrolateral glands distinct, forming a broken line, (4) ventral surface yellowish creamy-white with marble texture, (5) iris copper in the upper and silver in the lower fifth and (6) toe webbing basal and narrow lateral fringes. The male advertisement call of L. shangsiensis sp. nov. consists of one note per call, lacking an introductory note, with a mean dominant frequency of 5.5–6.5 kHz at the ambient temperature of 21.5ºC. Uncorrected sequence divergences between L. shangsiensis sp. nov. and all homologous DNA sequences available for the 16S rRNA gene were found to be > 5.9%. L. shangsiensis sp. nov. occurs in an evergreen forest in Shiwandashan National Nature Reserve located at an elevation of 450–550 m.

Two new species of the genusLeptobrachellaSmith, 1925,L.bijieJ. Wang, Y.L. Li, Y. Li, H.H. Chen & Y.Y. Wang,sp. nov.andL.purpuraventraJ. Wang, Y.L. Li, Y. Li, H.H. Chen & Y.Y. Wang,sp. nov., were described from northwestern Guizhou Province, China based on a combination of acoustic, molecular, and morphological data. The new discoveries bring the total number of this genus to 73, with 16 congeners recorded in China, and represent the second and third species of the genus reported from Guizhou Province.

Based on morphological and phylogenetic analyses (16S rRNA mtDNA), two new species of the genusLeptobrachellaare described from southern China, namelyL.yunkaiensisWang, Li, Lyu & Wang,sp. n.from Dawuling Forest Station of Guangdong Province andL.wuhuangmontisWang, Yang & Wang,sp. n.from Mt. Wuhuang of Guangxi Province. To date, the genusLeptobrachellacontains 68 species, among which 13 species are known from China. The descriptions of the two new species further emphasize that the species diversity of the genusLeptobrachellafrom China is still highly underestimated and requires further investigations.

We describe a tiny megophryid frog Leptobrachella juliandringi sp. nov. from Northeastern to Central Sarawak, Malaysian Borneo. This species is distributed in hilly and montane regions, and has long been confused with L. mjobergi. Based on the assessment using morphological and bioacoustic approaches, we conclude that the new species is distinct from all the other congeneric species including topotypic L. mjobergi from western Sarawak in possessing a series of elongate glands ventrolaterally on the trunk, broken reticulated marking on the ventrum, strap-like lineae masculinae, and an advertisement call with a chirp. The new species is also differentiated phylogenetically from the other congeners, and tends to form a group with true L. mjobergi but with a large genetic divergence.

Southeast Asia and southern China (SEA-SC) harbor a highly diverse and endemic flora and fauna that is under increasing threat. An understanding of the biogeographical history and drivers of this diversity is lacking, especially in some of the most diverse and threatened groups. The Asian leaf-litter frog genus Leptolalax Dubois 1980 is a forest-dependent genus distributed throughout SEA-SC, making it an ideal study group to examine specific biogeographic hypotheses. In addition, the diversity of this genus remains poorly understood, and the phylogenetic relationships among species of Leptolalax and closely related Leptobrachella Smith 1928 remain unclear. Herein, we evaluate species-level diversity based on 48 of the 53 described species from throughout the distribution of Leptolalax. Molecular analyses reveal many undescribed species, mostly in southern China and Indochina. Our well-resolved phylogeny based on multiple nuclear DNA markers shows that Leptolalax is not monophyletic with respect to Leptobrachella and, thus, we assign the former to being a junior synonym of the latter. Similarly, analyses reject monophyly of the two subgenera of Leptolalax. The diversification pattern of the group is complex, involving a high degree of sympatry and prevalence of microendemic species. Northern Sundaland (Borneo) and eastern Indochina (Vietnam) appear to have played pivotal roles as geographical centers of diversification, and paleoclimatic changes and tectonic movements seem to have driven the major divergence of clades. Analyses fail to reject an "upstream" colonization hypothesis, and, thus, the genus appears to have originated in Sundaland and then colonized mainland Asia. Our results reveal that both vicariance and dispersal are responsible for current distribution patterns in the genus.

The axial skeleton in most anuran families consists of or=7). Tadpoles from each genus are typically found in streams, where their extended caudal skeleton anchors muscles that facilitate tadpoles wiggling between plant debris and rocks or even burrowing into the stream bed. The extra centra of megophryids ossify differently in each genus. In Leptobrachella and Ophryophryne, the caudal centra ossify around the entire notochord, whereas in Megophrys and Xenophrys each develops from dorsal and ventral pairs of ossifications that expand to meet each other. The evolutionary loss of caudal centra, an apomorphic anuran trait, is reversed in larval megophryids and confirms that the machinery for caudal vertebral development has been retained in some modern anurans. A likely driving force in the reappearance of the trait in megophryids is the selective pressure associated with a riparian lifestyle.

Leptobrachella brevicrus Colour in life and preservation (stage 25) The following description is based on five specimens at Stage 25 whose total length ranged from 16.50 to 46.50 mm (Tables 2 and 3). The skin of the head, body and tail is translucent brown. Dorsally (Fig. 2 C), the head and body are uniformly brown with small brown melanocytes. The colour of the snout is darker coffee brown, getting slightly lighter behind the ocular region. During ontogenesis the brown pigmentation changes from a light yellowish brown in smaller tadpoles to darker brown in larger specimens. In lateral view (Fig. 2 A), the brown pigmentation of the trunk changes from brown to an iridescent bluish-white colour. The spiracle is translucent and therefore difficult to distinguish from the body wall in lateral view. The lateral line system is clearly visible against the background colour. The nares have a beige interior surrounded by darker coffee brown pigmentation. The oral funnel is translucent. The eyes are black, with dark grey pupils in larger specimens. The pigmentation of the tail is light yellowish brown and the myotomes and myosepta are visible. Stage n BL BH BS BW ED ES IND IOD LFH MTH The dorsal and ventral fins are both unpigmented and opaque, pale yellowish brown. The tip of the tail is translucent. In ventral view (Fig. 2 B), the skin has a bluish���white sheen which is particularly intense between the oral disk and the hyobranchial apparatus. The skin of the dorsum and tail is translucent. In lateral and ventral views, the reddish gills, the pale yellowish liver and the gut coil are clearly visible through the skin. In preserved specimens (Fig. 2 F) the red colour of the gills and the brown pigmentation of head, body and tail disappear. The skin is transparent with a slightly darker bluish sheen. Myotomes and myosepta become more prominent. The oral funnel becomes translucent. The vena caudalis ventralis and vena caudalis lateralis are visible in ventral and lateral views. The eyes are uniformly black. External morphological features. The body shape is vermiform, the head and trunk are elongated and slender (Fig. 2 A). The trunk and head are subcylindrical. The head and belly are slightly depressed. The anterior tip of the head is rounded in lateral and dorsal views. The nares are closer to the snout than to the eyes (Fig. 2 C). The nares are round and open antero���laterally with a short dorsal projection. The internarial distance is subequal to the interorbital distance (Table 2). The eyes are positioned dorsolaterally. They are small and sunken and do not protrude beyond the body wall, the iris is uniformly black. The spiracle is sinistral, long and conical in shape and has a pointed end (Fig. 2 C). The spiracular tube protrudes from the body wall and opens laterally. The opening of the spiracle is positioned at 48���52 % of the head���trunk length. The tail makes up 70 % of the total body length. The dorsal fin is low in the proximal half of the tail, only broadening slightly in the posterior half. The height of the ventral fin is constant in the proximal half of the tail and only increases slightly in the posterior half. The upper and lower fin each contributes 25 % to the maximum tail height. The tip of the tail is rounded. The muscles are visible through the transparent skin. The width of the base of the tail is 70���80 % of the maximal trunk width. The anal siphon is short and dextral. The subterminal, antero-ventrally protruding oral disc can be seen from the both ventral and lateral aspects. The width of the funnel-shaped oral disc makes up 80 % of the maximal width of the trunk (Fig. 2 A). The upper and lower lips each have deep medial emarginations (Fig. 2 E). The short and moderately pointed marginal papillae are arranged in a single row. Submarginal papillae are present on both sides between the oral orifice and the emarginations of the upper and lower lip and around the oral orifice. Below the submarginal papillae on the upper lip, two parallel parasagittal ridges emerge, run towards the upper jaw sheath and diverge laterally. On the lower lip a medial broad pad of thickened skin, parallel narrow ridges and isolated papillae are present. Keratodonts are absent. The beak is black and well keratinized with sharp serrations. The upper beak is arched in ventral view whereas the lower jaw is V-shaped. At Stage 39 the eyes protrude beyond the contour of the body. The oral disk and spiracle are reduced. All five toes are fully separated. Toe pads and subarticular tubercles are present. The tail is elongate and shows thick caudal muscles, the dorsal and ventral fins are not yet reduced., Published as part of Oberhummer, Evelyne, Barten, Catherin, Schweizer, Manuel, Das, Indraneil, Haas, Alexander & Hertwig, Stefan T., 2014, Description of the tadpoles of three rare species of megophryid frogs (Amphibia: Anura: Megophryidae) from Gunung Mulu, Sarawak, Malaysia, pp. 59-79 in Zootaxa 3835 (1) on pages 66-69, DOI: 10.11646/zootaxa.3835.1.3, http://zenodo.org/record/286717

The Leptolalax applebyi group of Asian leaf-litter frogs currently comprises four species of particularly small-bodied (

Tadpoles of the Megophryidae, a South Asian family of litter frogs, are unique among anurans by virtue of their expanded caudal skeletons, which include supernumerary vertebral centra. The number of these vertebrae varies widely within the family, with tadpoles of Leptobrachella having as many as 30 and Leptolalax only five. Vertebral morphology is also quite variable, ranging from complete, perichordal centra to fragmentary ossifications. This variation in the caudal osteology of larval megophryids, however, is not manifested in the adult morphology. Post-metamorphic litter frogs have a typical anuran axial skeleton, invariably comprising eight presacral vertebrae, a single sacral vertebra and, postsacrally, the urostyle. To resolve this incongruity between life phases and to determine the precise metamorphic fate of supernumerary caudal vertebrae in megophryids, we examined metamorphic specimens from the genera Leptobrachella, Leptolalax, Ophryophryne and Megophrys. In all four, the caudal larval skeleton undergoes massive reduction, leaving only the coccyx and hypochord untouched. Caudal centra are apparently degraded by osteoclasts, which have not previously been implicated in vertebral remodelling during anuran metamorphosis. In Megophrys and Ophryophryne metamorphs, presacral centra also undergo resorption, consistent with an epichordal mode of centrum formation. The conservation of megophryid adult axial osteology in the face of extensive larval skeletal diversity reveals the role of metamorphosis in constraining anuran morphology.

A new species of Leptolalax is described from Kinabalu National Park in western Sabah, Malaysian Borneo. The new species had been assigned to L. dringi, L. gracilis, or L. fritinniens in the past. It differs from all congeners, including these species, by a unique combination of morphological characters, including small body size, rounded snout, narrower interorbital than upper eyelid, basal toe webbing, smooth skin with tiny tubercles on dorsum and dorsal side of head, small pectoral glands, absence of supraaxillary glands and ventrolateral glandular ridges, spotted venter, advertisement call consisting of long series of 1–149 notes, each composed of three or four pulses, and dominant frequency at 6.90–7.35 kHz, without prominent frequency modulation.

Six species are recognized: from Borneo, L. baluensis SMITH, L. mjobergi SMITH, L. parva sp.n. and L. brevicrus sp. n.; from the Bunguran Islands,L. natunae(GUNTHER) andL. serasanae sp. n. Calls ofthe Bornean species are described.

In this paper the generic and subgeneric classification of the Pelobatidae Megophryinae is reexamined. Four genera are recognized in this subfamily. The first one, Megophrys Kuhl & Van Hasselt, 1822, includes two subgenera, the nominal one and Ophryophryne Boulenger, 1903, the latter being removed from the Bufonidae where several authors had placed it, and transferred into the Pelobatidae, as already suggested by Noble (1926) and by Liu & Hu (1962). For the second genus, the name Leptobrachella Smith, 1925 is revived in the place of Nesobia Van Kampen, 1923, the latter name being preoccupied by Nesobia Ancey, 1887 (Mollusca). The third genus is Lep tobrachium Tschudi, 1838, with two subgenera, the nominal one, of which Vibrissaphora Liu, 1945 is here considered to be synonymous, and a new subgenus, created for Leptobrachium gracile Gunther, 1872 and related species. The fourth genus, Scutiger Theobald, 1868, includes two subgenera, the nominal one and Oreolalax Myers & Leviton, 1962. The main diagnostic features of the genera and subgenera recognized here are given in tables., Dubois Alain. Notes sur la systématique et la répartition des Amphibiens Anoures de Chine et des régions avoisinantes. IV. Classification générique et subgénérique des Pelobatidae Megophryinae. In: Bulletin mensuel de la Société linnéenne de Lyon, 49ᵉ année, n°8, octobre 1980. pp. 469-482.