Bioacoustics of Leptobrachella murphyi

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: II<I<IV<III; tips of fingers rounded and slightly swollen; subarticular tubercles absent on fingers, inner metacarpal tubercle large and rounded, separated from much smaller outer metacarpal; supra-axillary glands oval. Hindlimb relatively long, tibio-tarsal articulation of adpressed limb reaching beyond eye, but not reaching snout; heels meeting when the tibias perpendicular to the body axis; tibia length less than half of snout-vent length (TIB/SVL=0.46); relative toe length: I<II<V<III<IV; toe tips rounded and slightly swollen; rudimentary webbing present between all five toes; wide lateral fringes present on all toes; subarticular tubercles indistinct under the base of I, II and III toe; distinct dermal ridges present under toes and uninterrupted at the articulations; moderate, oval inner metatarsal tubercle distinct, outer metatarsal tubercle absent ( Fig. 4 ). Dorsal skin relatively rough with small tubercles and short folds with reddish tips; ventral skin smooth; supra-axillary gland distinct and yellowish, located in axillary region dorsally from insertion of forelimb; pectoral gland small and indistinct; round femoral glands present and protuberant on rear of thigh, closer to knee than to vent; ventrolateral glands forming a distinct white line on flanks ( Fig. 4 ). Coloration 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 ).

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). Species Leptobrachella murphyi sp. nov. Holotype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Voucher Numbers KIZ 034039 KIZ 034159 KIZ 031199 KIZ 034183 KIZ 034225 KIZ 034173 KIZ 031200 KIZ 034182 KIZ 031168 KIZ 031198 KIZ 034166 KIZ 034158 KIZ 034038 KIZ 034165 Sex Male Male Male Male Male Male Male Male Male Male Female Female Female Female SVL 24.6 23.2 23.2 23.3 23.3 23.5 23.7 24.3 24.8 24.9 29.3 29.5 31.9 32.1 HDL 9.0 8.6 8.8 9.1 9.1 8.3 8.9 8.9 9.1 9.1 10.1 10.1 10.9 10.1 HDW 8.7 8.4 7.9 9.4 9.2 8.4 8.0 8.9 8.6 8.5 10.1 10.5 10.7 11.4 SNT 3.4 3.5 3.6 3.4 3.5 3.3 3.6 3.6 3.1 3.5 4.5 4.2 4.5 4.7 IND 2.4 2.2 2.2 2.1 2.1 2.3 2.4 2.1 2.4 2.4 2.8 2.6 3.0 2.8 IOD 3.2 3.1 3.1 2.4 2.7 2.8 3.2 2.8 3.1 2.7 3.4 3.4 3.6 3.8 UEW 2.3 2.3 2.4 2.1 2.3 2.2 2.0 2.3 2.3 2.0 2.9 2.8 3.0 3.1 EYE 3.3 3.4 3.1 3.4 3.6 3.5 3.2 3.6 3.1 3.3 4.0 4.1 4.0 4.2 TMP 1.9 2.0 1.5 2.1 2.0 2.0 1.5 2.1 1.7 1.8 2.4 2.3 2.4 3.1 TEY 1.2 1.2 0.8 1.0 1.1 1.0 0.7 1.2 1.0 1.2 1.2 1.2 1.4 1.4 NSL 1.3 1.4 1.5 1.6 1.6 1.5 1.3 1.6 1.4 1.4 2.0 1.7 1.9 1.7 NEL 2.0 1.9 1.8 1.8 1.6 1.7 2.2 1.6 2.0 2.0 2.5 2.3 2.6 2.7 FAL 6.4 5.9 5.1 6.3 6.6 6.4 5.4 6.7 6.1 5.7 7.3 7.6 7.5 8.1 ML 7.0 6.3 6.3 6.5 6.4 6.3 6.4 6.9 6.4 6.9 8.1 7.7 8.2 8.0 THL 12.2 10.9 11.4 11.6 11.4 11.1 11.2 12.7 12.5 12.2 14.7 14.5 14.9 15.3 TIB 11.4 11.5 11.6 10.6 11.2 10.3 12.2 10.8 12.6 12.7 15.1 14.7 15.4 15.3 TFOL 19.1 18.0 17.7 18.1 17.6 17.7 17.2 17.6 19.2 19.2 22.6 21.8 23.8 23.0 IMT 1.1 1.1 1.0 1.0 1.1 1.0 1.1 1.1 1.2 1.0 1.3 1.4 1.5 1.5 FIIIL 4.5 4.2 4.1 4.5 4.5 4.3 4.3 4.7 4.2 4.1 5.4 5.2 5.9 5.5 FEM 1.2 1.0 1.1 1.0 0.9 1.1 0.9 0.9 1.2 1.0 1.1 1.1 1.1 1.4 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). Species Male SVL (mm) Black spots on flanks Toes webbing Fringes on toes Ventral coloration Dorsal skin texture murphyi sp. nov. 23.2–24.9 Yes Rudimentary Wide Creamy white belly with small black spots on the margin Shagreened with reddish tubercles and folds aerea 25.1–28.9 No Rudimentary Wide Near immaculate creamy white, brown specking on margins Finely tuberculate alpina 24.0–26.4 Yes Rudimentary Wide in males Creamy-white with dark spots Relatively smooth, some with small warts applebyi 19.6–22.3 Yes Rudimentary Absent Reddish brown with white speckling Smooth ardens 21.3–24.7 Yes Absent Absent Reddish brown with white speckling Smooth-finely shagreened aspera 22.4 Yes Rudimentary Narrow Creamy white with distinct dark patches on chest and abdomen Rough with dense conical granules, tubercles, and glandular folds bashaensis 22.9–25.6 Yes Rudimentary Narrow Creamy-white chest and off-white belly with irregular black spots slightly shagreened with small tubercles and irregular brown stripes bidoupensis 18.5–25.4 Yes Rudimentary Narrow Reddish brown with white speckling Smooth bijie 29.0–30.4 Yes Rudimentary Narrow White with distinct nebulous greyish speckling on chest and ventrolateral flanks Shagreened and granular botsfordi 29.1–32.6 No Rudimentary Narrow Reddish brown with white speckling Shagreened bourreti 28.0–36.2 Yes Rudimentary Narrow Creamy white Relatively smooth, some with small warts chishuiensis 30.8–33.4 Yes Rudimentary Narrow White with distinct nebulous greyish speckling on chest and ventrolateral flanks Shagreened and granular crocea 22.2–27.3 No Rudimentary Absent Bright orange Highly tuberculate dorsospina 28.7–30.5 Yes Rudimentary Narrow Greyish white with black spots and orange pigmenta- tions Rough with dense conical granules, tubercles, glandular folds, and conical spines eos 33.1–34.7 No Rudimentary Wide Creamy white Shagreened ......continued on the next page Species Male SVL (mm) Black spots on flanks Toes webbing Fringes on toes Ventral coloration Dorsal skin texture feii 21.5–22.8 Yes Rudimentary Narrow Creamy white with black blotches Shagreened with small tubercles and ridge firthi 26.4–29.2 No Rudimentary Wide in males Creamy white Shagreened with fine tubercles flaviglandulosa 23.0–27.0 Yes Rudimentary Narrow Whitish, black speckling on margins Shagreened with yellowish-brown tubercles fuliginosa 28.2–30.0 Yes Rudimentary narrow White with brown dusting Nearly smooth, few tubercles graminicola 23.1–24.6 In some individuals Rudimentary Wide White with very dark blackish brown spots Smooth, with many tubercles and lacking dermal ridges isos 23.7–27.9 No Rudimentary Wide in males Creamy white with white dusting on margins Mostly smooth, females more tuberculate jinshaensis 29.7–31.2 Yes Absent Narrow Cream yellow with purple speckling, and on flanks presence of distinct nebulous greyish speckling Shagreened, some of the granules forming longitudinal short skin ridges kalonensis 25.8–30.6 Yes Absent Absent Pale, speckled brown Smooth khasiorum 24.5–27.3 Yes Rudimentary Wide Creamy white Isolated, scattered tubercles lateralis 26.9–28.3 Yes Rudimentary Absent Creamy white Roughly granular laui 24.8–26.7 Yes Rudimentary Wide Creamy white with dark brown dusting on margins Round granular tubercles liui 23.0–28.7 Yes Rudimentary Wide Creamy white with dark brown spots on chest and margins Round granular tubercles with glandular folds macrops 28.0–29.3 Yes Rudimentary Absent Greyish-violet with white speckling Roughly granular with larger tubercles maculosa 24.2–26.6 Yes Absent Absent Brown, less white speckling Mostly smooth mangshanensis 22.2–27.8 Yes Rudimentary Narrow Throat grey-white and belly creamy white, scattered with white speckles Smooth with orange tubercles and dark brown stripes maoershanensis 25.2–30.4 Yes Rudimentary Narrow Creamy white chest and belly with irregular black spots Longitudinal folds melica 19.5–22.7 Yes Rudimentary Absent Reddish brown with white speckling Smooth ......continued on the next page Species Male SVL (mm) Black spots on flanks Toes webbing Fringes on toes Ventral coloration Dorsal skin texture minima 25.7–31.4 Yes Rudimentary Absent Creamy white Smooth nahangensis 40.8 Yes Rudimentary Absent Creamy white with light specking on throat and chest Smooth neangi 35.4–36.3 (Females) Yes Absent Absent Belly transparent, immaculate purplish gray Dorsal skin with small, irregular bumps and ridges niveimontis 22.5–23.6 Yes Rudimentary Narrow Marbled with black speckling Relatively smooth with small tubercles nokrekensis 26.0–33.0 Yes Rudimentary Unknown Creamy white Tubercles and longitudinal folds nyx 26.7–32.6 Yes but indistinct Rudimentary Absent Creamy white with white with brown margins Rounded tubercles oshanensis 26.6–30.7 Yes Absent Absent Whitish with no markings or only small, light grey spots Smooth with few glandular ridges pallida 24.5–27.7 No Absent Absent Reddish brown with white speckling Tuberculate pelodytoides 27.5–32.3 Yes One third webbing Narrow Whitish Mostly smooth with smooth warts petrops 23.6–27.6 No Absent Narrow Immaculate creamy white Highly tuberculate pluvialis 21.3–22.3 Yes Rudimentary Absent Dirty white with dark brown marbling Smooth, flattened tubercles on flanks puhoatensis 24.2–28.1 Yes Rudimentary Narrow Reddish brown with white dusting Longitudinal skin ridges purpuraventra 27.3–29.8 Yes Rudimentary Narrow Grey purple with distinct nebulous greyish speckling on chest and ventrolateral flanks Shagreened and granular purpurus 25.0–27.5 Yes Rudimentary Wide Dull white with indistinct grey dusting Shagreen with small tubercles pyrrhops 30.8–34.3 Yes Rudimentary Absent Reddish brown with white speckling Slightly shagreened rowleyae 23.4–25.4 Yes Absent Absent Pinkish milk-white to light brown with white speckles Smooth with numerous tiny tubercles shiwandashanensis 26.8–29.7 Yes Absent Absent Creamy white with brown spots on the lateral margin Rough with numerous tubercles ......continued on the next page Species Male SVL (mm) Black spots on flanks Toes webbing Fringes on toes Ventral coloration Dorsal skin texture suiyangensis 28.7–29.7 Yes Rudimentary Narrow Yellowish creamy-white with marble texture chest and belly or with irregular light brown speckling Shagreen with small granules sungi 48.3–52.7 No or small Wide Narrow White Granular tadungensis 23.3–28.2 Yes Absent Absent Reddish brown with white speckling Smooth tamdil 32.3 Yes Wide Wide White Weakly tuberculate tengchongensis 23.9–26.0 Yes Rudimentary Narrow White with dark brown blotches Shagreened with small tubercles tuberosa 24.4–29.5 No Rudimentary Absent White with small grey spots/streaks Highly tuberculate ventripunctata 25.5–28.0 Yes Rudimentary Absent or narrow Chest and belly with dark brown spots Longitudinal skin ridges wuhuangmontis 25.6–30.0 Yes Rudimentary Narrow Greyish white mixed by tiny white and black dots Rough with dense conical tubercles wulingensis 24.5–32.8 Small to moderate Rudimentary Narrow Creamy white, with distinct or indistinct brown speck- ling at margins Shagreened with sparse large warts, sometimes with longitudinal ridges yeae 25.8–32.6 Yes Rudimentary Narrow Cream white with small brown specking on sides and upper abdomen Relatively smooth with tiny granules yingjiangensis 25.7–27.6 Yes Rudimentary Wide Creamy white with dark brown flecks on chest and margins Shagreened with small tubercles yunkaiensis 25.9–29.3 Yes Rudimentary Wide Belly pink with distinct or indistinct speckling Shagreened with short skin ridges and warts zhangyapingi 45.8–52.5 Yes Rudimentary Wide Near immaculate white Mostly smooth with distinct tubercles (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 invariably single-note call with irregular intervals. In addition, the dominant frequency of 4.5–4.7 kHz (at 15.0 °C) further distinguishes the call of L . murphyi sp. nov. from that of the higher frequency calls of L. aereus (6.2–7.9 kHz at 22.4–25.7 °C), L. yingjiangensis (5.7–5.9 kHz at 19 °C), L. bijie (4.8–5.1 kHz at 18.6–19.3 °C), L. shangsiensis (5.5–6.5 kHz at 21.5 °C), L. isos (5.9–6.2 kHz at 22.4–22.8 °C), L. puhoatensis (4.9–5.6 kHz at 22.3–25.8 °C) and L. ventripunctatus (6.1–6.4 kHz at 15 °C), and the lower frequency calls of L. applebyi (4.0–4.3 kHz at 21.5 °C), L. ardens (3.1–3.4 kHz at 21.4–24.7 °C), L. bidoupensis (1.9–3.8 kHz at 19–21 °C), L. botsfordi (2.6–3.2 kHz at 14 °C), L. croceus (2.6–3.0 kHz at 21.6–25.1 °C), L. fuliginosus (2.1–2.8 kHz at 19.3–19.6 °C), L. kalonensis (2.8 kHz at 26.4 °C), L. maculosus (2.7–2.8 kHz at 23.3–24.1 °C), L. melicus (2.6–4.0 kHz at 26.1–26.2 °C), L. pallidus (2.4–2.7 kHz at 14.0–21.4 °C), L. purpurus (4.3–4.5 kHz at 15 °C), L. pyrrhops (1.91–2.2 kHz at 25 °C), L. rowleyae (3.3–3.5 kHz at 21.5 °C), L. tadungensis (2.6–3.1 kHz at 12.9–22.3 °C) and L. tuberosus (2.6–2.8 kHz at 22.5–24.5 °C).