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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"]}

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

Chen, Weicai, Liao, Xiaowen, Zhou, Shichu, Mo, Yunming, Huang, Yong (2018): Rediscovery of Rhacophorus yaoshanensis and Theloderma kwangsiensis at their type localities after five decades. Zootaxa 4379 (4): 484-496, DOI: https://doi.org/10.11646/zootaxa.4379.4.2

Theloderma kwangsiensis Liu & Hu, 1962 Type locality. Dayaoshan Ranges, Jinxiu County, Guangxi, China. Specimens examined (n=4). NHMG201504001, adult male, was collected at the type locality, the Dayaoshan Ranges, Guangxi, China (24.168397�� N, 110.244343�� E, 1150 m asl). The other three individuals (NHMG201608026, adult male; NHMG20161003, adult male, and NHMG20161101, adult female) were collected at Shiwandashan National Nature Reserve, Guangxi, China (21.844043�� N, 107.891647�� E, 532 m asl). NHMG1504001, NHMG201608026 and NHMG20161003 were found in PVC buckets (diameter 25 cm, height 20 cm) which were used to monitor amphibians. NHMG20161101 was found on a tree approximately 0.5 m above the ground in the evergreen forest. Description. Body dorsoventrally compressed; head length less than head width, head very strongly depressed; snout pointed in dorsal view and profile, projecting beyond margin of the lower jaw; canthus rostral distinct, loreal region sloping; nostrils oval, oblique, nearly reaching the tip of the snout; eye diameter less than snout length; interorbital region concave; interorbital distance slightly less than eye diameter; internarial distance less than eye diameter; pupil horizontal; tympanum distinct, rounded, less than eye diameter, slightly convex relative to skin of temporal region; vomerine teeth in two small oblique groups nearly linking the anterior of the choana; tongue pear-shaped with notch posterior; vocal sac absent. (Table 3; Fig. 5 A���D). Forelimbs moderately robust, relative length of fingers IColoration of T. kwangsiensis in life. The dorsal surface is grass-green with irregular, army-green blotches which consist of large irregular warts studded with small dark red granules. The venter covers cream yellow granules with irregular pale brown spots. The pupil is oval, and the iris is black with cream-yellow reticulations throughout. Ecology. Three individuals of T. kwangsiensis were found in PVC buckets (artificial refugia for monitoring amphibians, diameter= 25 cm, height= 20 cm) on 25 April 2015 (NHMG 201504001), on 25 August 2016 (NHMG 201608026) and on 14 October 2016 (NHMG 20161003). One individual (NHMG 20161101) was found on a tree approximately 0.5 m above the ground in evergreen forest (Fig. 7). Distribution. Currently, we have confirmed that T. kwangsiensis occurs in the Dayaoshan Ranges and Shiwandashan National Nature Reserve. Molecular analyses. The genetic variation between the Dayaoshan Ranges individual and Shiwandashan National Nature Reserve individuals ranged from 0.0���0.6% based on the 16S and 12S DNA fragments sequenced (~2000 bp). Theloderma kwangsiensis nested within T. corticale with high support, and the phylogenetic tree was consistent with Poyarkov et al. (2015), Huang et al. (2017), and Hou et al (2017). The genetic distance between T. kwangsiensis and T. corticale ranged from 0.0���0.6%., Published as part of Chen, Weicai, Liao, Xiaowen, Zhou, Shichu, Mo, Yunming & Huang, Yong, 2018, Rediscovery of Rhacophorus yaoshanensis and Theloderma kwangsiensis at their type localities after five decades, pp. 484-496 in Zootaxa 4379 (4) on pages 492-495, DOI: 10.11646/zootaxa.4379.4.2, http://zenodo.org/record/1174988, {"references":["Liu, C. C. & Hu S. Q. (1962) A herpetological report of Kwangsi. Acta Zoologica Sinica, 14 (Supplement), 73 - 104.","Poyarkov, N. A. J., Orlov, N. L., Moiseeva, A. V., Pawangkhanant, P., Ruangsuwan, T., Vassilieva, A. B., Galoyan, E. A., Nguyen, T. T. & Gogoleva, S. S. (2015) Sorting out moss frogs: mtDNA data on taxonomic diversity and phylogenetic relationships of the Indochinese species of the genus Theloderma (Anura, Rhacophoridae). Russian Journal of Herpetology, 22, 241 - 280.","Huang, H., Chen, Z., wei, Z., Bu, R. & wu, Z. (2017) DNA barcoding revises a misidentification on mossy frog: new record and distribution extension of Theloderma corticale, Boulenger, 1903 (Amphibia: Anura: Rhacophoridae). Mitochondrial DNA, Part A, DNA Mapping, Sequencing, and Analysis, 29 (2), 273 - 280."]}

Mo, Yunming, Zhang, Wei, Zhou, Shichu, Chen, Tianbo, Tang, Huaxing, Meng, Yuanjun, Chen, Weicai (2013): A new species of Kaloula (Amphibia: Anura: Microhylidae) from southern Guangxi, China. Zootaxa 3710 (2): 165-178, DOI: http://dx.doi.org/10.11646/zootaxa.3710.2.3

Mo, Yunming, Zhang, Wei, Luo, Yu, Zhou, Shichu, Chen, Weicai (2013): A new species of the genus Gracixalus (Amphibia: Anura: Rhacophoridae) from Southern Guangxi, China. Zootaxa 3616 (1): 61-72, DOI: http://dx.doi.org/10.11646/zootaxa.3616.1.5

Kaloula nonggangensis sp. nov. Holotype. NHMG 1106036, adult male, from the Nonggang National Nature Reserve, Southern Guangxi Province, China (22.4522 �� N, 106.9354 �� E; altitude: 186 m a.s.l.), collected on June 28, 2011 by Weicai Chen, Yunming Mo (Figure 2 A, B, C). Paratypes. NHMG 1106030 ��� 35, NHMG 1106037 ��� 41, adult males collected at the same place of the locality by Weicai Chen and Yunming Mo on June 29, 2011. NHMG 1108035, adult female, NHMG 1108036 ��� 41 adult males, collected at the near locality of the holotype (22.4809 �� N, 106.9017 �� E; altitude: 167 m a.s.l.) by Yunming Mo, Wei Liao and Zhuqiu Song on August 11, 2011. Diagnosis. Assigned to the genus Kaloula on the basis of the following: medium size (41.4���52.7 mm in 18 adult male, 52.2 mm in one adult female), smooth or slightly rough olive dorsum with irregular dark-green marks and brown spots (Figure 2 A, B); tips of the fingers dilate and truncated (Figure 3 A); nearly full webbing on toes in males and reduced webbing in females (Figure 3 B); two side protuberant osseous tubercles on the upper surface of the tips of fingers in male (Figure 3 C); chest beige with small lemon-colored spots in male (Figure 2 C); ventral epidermal adhesive gland occupies chest and venter; larvae lacking a keratinized jaw sheath and labial teeth. K. nonggangensis sp. nov. is distinguished from it congeners by a combination of (1) medium size (SVL ranging 41.4���52.7 mm in males), (2) smooth or slightly rough olive dorsum with irregular dark-green marks and brown spots, (3) tips of the fingers dilate and truncated, (4) chest beige with small lemon-colored spots in male, and (5) male with two side protuberant osseous tubercles on the upper surface of the tips of fingers. Description of holotype. Adult male; SVL 47.5 mm; habitus rotund, body rounded; head proportionally small, length 25 % of SVL, 76 % of head width; snout slightly pointed beyond lower jaw, its tip round in dorsal aspect and in lateral aspect; eyes protruding laterally beyond silhouette of head in dorsal aspect, protruding very markedly beyond dorsal surface of head in lateral aspect; pupil circular; interorbital region flat with some small tubercles; snout less than half of the head length; upper eyelid width slightly smaller than interorbital distance, and much smaller than snout length; eye diameter (4.63 mm) almost equal to snout length (4.64 mm), longer than interorbital distance (3.8 mm) and eye-narial distance (2.1 mm); canthus rostralis indistinct; loreal region sloping, moderately concave; nostrils barely protuberant, very close to tip of snout (eye-nostril distance longer than distance from nostril to tip of snout); internarial region slightly concave; tympanum hidden and indistinct; tympanic fold prominent, extending from posterior corner of eye to supra-axillary region; tongue oval without posterior notch; choanae elongate oval and small, situated at anterolateral edge of palate, separated by a distance about two times of their diameter; dentigerous process of vomer in posterior edge of choanae evident, almost touching in the middle line but vomerine teeth apparently absent; vocal slits large, just posterior to rictus. Forelimbs. Arms short, forearms not hypertrophied; hand relatively large (HAL 14.3 mm); forearm and hand length (LAHL 23.3 mm) almost as long as half of SVL (47.5 mm); fingers disks obviously dilated and tips truncated; relative length of fingers I Hindlimbs. Hind limbs relatively short and brawny; heels not overlapping for long distance when legs are folded at right angle to body, and when appressed to body, tibiotarsal articulation reaching the shoulder; tibia 2.7 times longer (TL 19.7 mm) than wide (TW 7.4 mm), and shorter than foot length (FOL 21.3 mm), about 42 % of SVL. Relative length of toes: I Skin. Skin of dorsal surfaces of body, head, and limbs smooth; ventral surfaces of throat, trunk, and limbs smooth; loose skin overlying median subgular vocal sac forms slight sternal fold; ventral epidermal adhesive gland occupies sternal region and venter. Color of holotype in life. Dorsal parts of head and dorsum, flank, forelimb, thigh, tibia and foot light olivegreen with dark moss-green marbling; loreal region, tympanic region and tympanum slightly dark olive-green; dorsal parts of two sides shoulder light olive-green; lower part of flanks with some lemon spots; lower jaw, throat, margin of throat and chest beige with some small lemon spots; belly, ventral part of forelimbs and hind limbs creamy white. Color of holotype in preservative. Dorsum brown with dark marbling. Ventral surface of chest, throat, belly, interior portions of arms and thighs cream color. Measurements. Holotype: SVL 47.5, HL 11.7, HW 15.2, SL 4.6, IND 2.8, IOD 4.1, UEW 2.7, ED 4.6, LAHL 23.3, HAL 14.3, HLL 62.7, TL 19.7, TW 7.1, FOL 21.3. Etymology. The species epithet is a Latin adjective referring to the locality in which the new species was collected. The suggested English name is the Nonggang narrow-mouthed frog. Tadpole. Six tadpoles (Voucher No.: NHMG_T 20110801, NHMG_T 20120901 ���05) from stages 36 to 42 sensu Gosner (1960) were collected from the type locality of K. nonggangensis sp. nov. Morphological measurements and characteristics were examined following Altig & McDiarmid (1999). The 16 S rRNA sequence (~ 540 bp) of a specimen (Voucher No.: NHMG_T 20120901) is 99 % identical to a reference sequence of paratype (Voucher No.: NHMG 1108036). Measurements of one tadpole at developmental stage 38: total length 37.5 mm, head-body length 0.54 time of tail length; eyes lateral, small, and visible from ventral view. In dorsal view, body nearly rectangle, snout broadly rounded. Interpupilar distance 0.84 maximum body width, internarial distance 0.17 of interpupilar distance. Width of oral apparatus 0.27 head-body length; lips not expanded; lacking labial teeth, horny beak and papillae. Nostrils closed at stage 38. Spiracle median, opening slightly anterior the end of body, free portion with an arched membrane, the inner wall attached to body wall; anal tube median and elongate, and the inner wall fused to ventral fin. The base of tail musculature strong, its height 0.35 of tail height; maximal tail height located at the proximal 1 / 2 of the tail, caudal muscles tapering gradually. Dorsal fin originates at the tail bodyjunction and the ventral fin originates at the ventral terminus of the body. The dorsal fin is nearly equal to the ventral fin in height. Dorsal fin and ventral fin have some yolk-yellow spots. Especially, dorsal fin near the tail body-junction has a yolk-yellow line. Tail tip broadly rounded. Head and body brownish with some faint yellow pigments (Figure 2 G, H). Variation. Individuals of the type series are generally similar in morphology. The female (NHMG 1108035) is two-third webbed (webbing formula, I 2 - ��� 2 + II 2 - ��� 3 III 2 ��� 3 IV 3 + ��� 2 - V). Two specimens (NHMG 1108035 ��� 36) have some small spinous tubercles in the anal region. Five specimens (NHMG1106033, 1108035���36, 1108038, 1108040) have two subarticular tubercles on the third toe, and the remaining specimens have three subarticular tubercles on the third toe. Four individuals (NHMG 1106040 ���42, 1108037) have a slightly rough dorsum scattered with a few small rounded tubercles. The tympanic fold is indistinct in six individuals (NHMG1106036, 1108037��� 39, 1108040��� 41). All males have two side protuberant osseous tubercles on the upper surface of the tips of fingers. Between two side protuberant osseous tubercles, four individuals have small tubercles ranging from 1 to 4 (NHMG 1106031: L II (left finger II), 3; NHMG 1106033: L II, 4; NHMG 1108036: R I (right finger I), 2; NHMG 1108039: L I, 1; R III, 1), which size is about 1 / 5 of two side protuberant osseous tubercles. Female also have two side protuberant osseous tubercles but which size is only 1 / 2 that of the male. In preservative, seven specimens (NHMG 1106031 ���34, 1106035, 1106039, 1108040) have a gray dorsum, and the remaining specimens are dark brown. Male secondary sexual characters. Nuptial pad absent; external subgular vocal sac present; throat with some lemon spots; chest and belly with epidermal adhesive gland. Ecology. K. nonggangensis sp. nov. were observed in primary or secondary karst evergreen forest, and cultivated fields near the forest. Presently, K. nonggangensis sp. nov. is only known from the Nonggang National Nature Reserve. Sites where we observed the species ranged from 150���200 m elevation. On June 29 and August 11, 2011, the species was observed to gather in the temporary plash after rainstorm. Sequence divergence. Uncorrected sequence divergences between K. nonggangensis sp. nov. sequences and all homologous sequences available on the genus Kaloula are listed in Table 2. The uncorrected p -distance genetic distance between K. nonggangensis sp. nov. and K. verrucosa tissues examined was 1.4���1.6 %, while that between K. nonggangensis sp. nov. and other species was> 1.8 % (Table 2). Preliminary hypothesis of phylogenetic relationship. Among 15 Kaloula species (Frost 2013), only 9 species' sequences (including K. baleata, K. borealis, K. conjuncta, K. mediolineata, K. picta, K. pulchra, K. rugifera, K. taprobanica, K. verrucosa) are available from GenBank. Based upon our preliminary molecular data, the genus Kaloula formed a monophyletic group with well-supported values (BP= 98; BBP=1.00), and was divided three clades (Figure 4). Clade A consists of Kaloula taprobanica Parker, 1934; Clade B consists of K. borealis, K. rugifera, K. verrucoca, and K. nonggangensis sp. nov. and Clade C consists of K. baleata van & Muller, 1836, K. conjuncta Peters, 1863, K. mediolineata Smith, 1917, K. picta Dum��ril & Bibron, 1841, and K. pulchra Gray, 1831. Comparisons: K. nonggangensis sp. nov. differs from all other species of Kaloula by having smooth or slightly rough dorsum without rough tubercles, finger tips widely expanded and truncated, two side protuberant osseous tubercles on the upper surface of the tips of fingers in male, throat and chest with some small lemon spots in male, and larvae with some yolk-yellow spots in body and tail. K. nonggangensis sp. nov. is most similar in appearance to K. borealis, K. rugifera and K. verrucosa (Figure 2). However, K. nonggangensis sp. nov. differs from K. borealis by having widely expanded terminal digital disks (vs. lacking expanded terminal digital disks in the latter); by two side protuberant osseous tubercles on the upper surface of the tips of fingers in male (without osseous tubercles in the latter, Figure 3 D); by having nearly full webbing (vs. 1 / 2 webbing in the latter); by throat and chest beige with some small lemon spots (vs. distinct black spots in the throat area in the latter) (Fei et al. 2009). Furthermore, in life, the tadpole of K. borealis has dark brown dorsal body and tail, without pigment and venter white or hoar (vs. the tadpole of K. nonggangensis sp. nov. has brownish head and body with some faint yellow pigments and dorsal fin near the tail body-junction has a yolk-yellow line) (Fei et al. 2009; Zhou et al. 2011). K. nonggangensis sp. nov. differs from K. rugifera by having a relatively bigger body size (SVL 41.4���52.7 mm in male vs. 35.5 ���43.0 mm in male for K. rugifera) (Table 3). It can be further distinguished from K. rugifera by finger tips widely expanded and truncated (slightly expanded in K. rugifera, Figure 3 F); by two side protuberant osseous tubercles on the upper surface of the tips of fingers in male (having two clusters of osseous tubercles instead of two side protuberant osseous tubercles in the latter) (Figure 3 F); by throat and chest with some small lemon spots in male (vs. cream without pigments); by larvae with some yolk-yellow spots in body and tail (vs. dorsum and tail dark brown, no pigment) (Fei et al. 2009; Zhou et al. 2011). K. nonggangensis sp. nov. also differs from K. verrucosa by smooth or slightly rough dorsum without rough tubercles (rough dorsal skin and tubercles present in K. verrucosa) (Figure 2 F); by finger tips widely expanded and truncated (non-expansion of terminal digital disks in the latter, Figure 3 E); by two side protuberant osseous tubercles on the upper surface of the tips of fingers in male (4���6 free osseous tubercles in the latter) (Figure 3 E); by larvae with some yolk-yellow spots in body and tail (vs. dorsum and tail dark brown with black pigment in the latter) (Fei et al. 2009). The morphological differences in the shape of terminal disks (Figure 3) have been assessed in large numbers of specimens in the studies of 83 for K. borealis, 100 for K. verrucosa and 67 for K. rugifera, partly in specimens from different localities (Kunming and Dali in K. verrucosa) (Appendix I, Fei et al. 2009). K. nonggangensis sp. nov. differs from K. aureata Nutphand, 1989, by having an olive dorsum without lateral bands (vs. bright yellow in median dorsum, and yellowish orange lateral bands along the back) (Pauwels & Ch��rot, 2006). K. nonggangensis sp. nov. differs from K. baleata by having a ventral adhesive gland in males (vs. venter without epidermal adhesive gland); by having olive dorsum without rough tubercles (vs. brownish dorsum with rough tubercles, and a bright yellow spots near axilla in the latter) (Pauwels et al. 1999). K. nonggangensis sp. nov. differs from K. conjuncta, K. kalingensis, K. kokacii and K. walteri by a larger body size (SVL 41.4���52.7 mm in males vs. 26.6���30.1 mm in K. conjuncta, 24.2���32.9 mm in K. kalingensis, 37.0��� 39.1 mm in K. kokacii, 24.5���31.5 mm in K. walteri (Diesmos et al. 2002). It also differs from K. conjuncta by the absence of rough dorsal tubercles (vs. tubercles distributed over the entire dorsum); by having olive dorsum (vs. brownish dorsum) (Diesmos et al. 2002). It also differs from K. kalingensis and, K. kokacii by its ventral adhesive gland in males (vs. venter without epidermal adhesive gland) (Brown et al. 2000; Diesmos et al. 2002). It also differs from K. walteri by the presence of a round, small, and pointed outer metatarsal tubercle (vs. absence, or presence of only a faint, very reduced outer metatarsal tubercle); by the presence of three subarticular tubercles on the fourth toe (vs. two); by having widely expanded terminal digital disks (vs. lacking expanded terminal digital disks in the latter) (Diesmos et al. 2002). K. nonggangensis sp. nov. clearly differs from K. assamensis, K. mediolineata, K. pulchra and K. taprobanica by absence (vs. presence) of dorsolateral bands. It also differs from K. assamensis by its inner metatarsal tubercle smaller (vs. larger) than first toe; lacking a vertebral stripe (vs. presence of a dark edged lemon vertebral stripe) (Das et al. 2004; Nath et al. 2011). It also differs from K. mediolineata by its finger tips widely expanded (vs. non expanded) and truncated (Diesmos et al. 2002). It also differs from K. pulchra by its smaller body size (SVL 41.4��� 52.7 mm vs. 55.0���77.0 mm in K. pulchra (Fei et al. 2009); snout length about 1 / 3 head length (vs. half of the head length; by smooth venter (vs. granular venter) (Fei et al. 2009). It can be further distinguished from K. taprobanica by lacking (vs. having) large irregular shaped markings of yellow-red color in dorsum median area (Dutta & Manamendra-Arachchi 1996). K. nonggangensis sp. nov. differs from K. picta and K. rigida by absence (vs. presence) of supernumerary tubercles at the base of each digit of the manus. It also differs from K. picta by its inner metatarsal tubercle shorter (vs. equal to or longer) than first toe (Diesmos et al. 2002). Finally, K. macrocephala (originally treated as a synonym of K. pulchra) differs from K. nonggangensis sp. nov. by having indistinct dorsolateral bands and mid-dorsum covered by large-sized irregular patches (vs. clearly absent dorsolateral bands in new species) (Bourret 1942; Ohler 2003; Pauwels & Ch��rot 2006). In addition, genetically, except for K. borealis, K. rugifera and K. verrucosa, the uncorrected sequence divergences between K. nonggangensis sp. nov. 16 S rRNA sequences and all homologous sequences available on GenBank (Table 2) were greater than 3 %, a value usually representing differentiation at the species level in frogs (Vences et al. 2005; Fouquet et al. 2007). Discussion Fei et al. (2009) divided Chinese Kaloula species into two groups: the K. pulchra group (consisting of K. pulchra) and the K. verrucosa group (consisting of K. borealis, K. rugifera and K. verrucosa), being consistent with our phylogenetic trees. K. nonggangensis sp. nov. embedded within the K. verrucosa group. K. nonggangensis sp. nov., K. borealis, K. rugifera and K. verrucosa formed a monophyletic group with high supported values (Figure 4). Preliminary molecular data indicated that K. nonggangensis sp. nov. and K. borealis were sister species. However, due to weak inter-nodes support values (K. verrucosa group are lower than the values usually representing differentiation at the species level in frogs (Vences et al. 2005; Fouquet et al. 2007) (Table 2). According to the results of these authors, in most cases species are differentiated by uncorrected p -distances in 16 S rRNA of 3 % or higher. If the criterion was strictly applied in this group, then K. nonggangensis sp. nov., K. borealis, K. ru, Published as part of Mo, Yunming, Zhang, Wei, Zhou, Shichu, Chen, Tianbo, Tang, Huaxing, Meng, Yuanjun & Chen, Weicai, 2013, A new species of Kaloula (Amphibia: Anura: Microhylidae) from southern Guangxi, China, pp. 165-178 in Zootaxa 3710 (2) on pages 168-175, DOI: 10.11646/zootaxa.3710.2.3, http://zenodo.org/record/216543

Gracixalus nonggangensis sp. nov. (Fig. 2) Holotype. NHMG 200809044 (Fig. 2), adult male, from Nonggang National Nature Reserve, Longzhou country, Guangxi, China, near the Sino-Vietnam border (22.5233 �� N, 106.9523 �� E, alt. 216 m a.s.l) (Fig. 1 A), collected by Yunming Mo, Wei Zhang and Shichu Zhou on 7 September 2008. Paratypes. NHMG 200809043, adult male collected near locality of the holotype (22.4567 �� N, 106.9742 �� E, alt. 432 m a.s.l) on 7 September 2008; NHMG 20091001, NHMG 20091003, NHMG 20091009 ��� 10, adult males, and NHMG 20091002, adult female, from the same site as the holotype, collected by Yunming Mo, Wei Zhang and Shichu Zhou on 14 October 2009. NHMG 201005008 and NHMG 111024, adult females, from the same site as the holotype, collected by Yunming Mo, Weicai Chen and Wei Zhang on 7 May 2010, and 14 October 2011. NHMG 1005046, adult male, from Longrui station of Nonggang National Nature Reserve, Ningming county (22.2463 �� N, 107.0729 �� E, alt. 252 m a.s.l), collected by Yunming Mo, Wei Liao and Zhuqiu Song on 10 May 2010. NHMG 111025, adult male, from the same site as the holotype, collected by Yunming Mo, Weicai Chen and Wei Zhang on 14 October 2011. Diagnosis. The new species is assigned to the genus Gracixalus by the tips of digits expanded into large disks, vomerine teeth absent, horizontal pupil, two outer palmar tubercles, tibiotarsal articulation reaching the tip of snout. Gracixalus nonggangensis sp. nov. can be distinguished from all other species of Gracixalus by the following combination of characters: medium size (SVL ranging from 29.9���35.3 mm for males, 33.6 ���38.0 mm for female); vomerine teeth absent; distinct tympanum as wide as the disc of finger III; lower part of tympanum with many small tubercles; tibiotarsal articulation reaching the tip of snout; dermal fringes along outer side of forearm, tibia and tarsus absent; smooth, yellowish-olive dorsum in life, with a wide dark-green irregular mark; throat, chest and belly white with light grey-blue tint and brown marbling in life; dark green, broad transverse stripes on limbs: 2 on the forearm, 3 on the thigh and tibia; finger webbing absent, toes one-third webbed; male with internal subgular vocal sac. Description of holotype. Adult male (SVL 30.0 mm), body moderately elongate; head slightly longer than wide (HL 12.0 mm; HW 11.2 mm; MN 10.0 mm; MFE 7.8 mm; MBE 3.4 mm); snout rounded in dorsal view and slightly protruding, SL (4.7 mm) longer than horizontal diameter of eye (EL 3.8 mm); canthal edges rounded, loreal region oblique, slightly concave; nostrils oval, markedly protuberant, directed laterally, closer to tip of snout (NS 1.6 mm) than to eye (EN3.0 mm); internasal distance (IN 2.9 mm) as wide as upper eyelid (UEW 2.8 mm), less than interorbital space (IUE 3.7 mm); internarial space concave; distance between front of eyes (IFE 5.9 mm) about 60 % distance between back of eyes (IBE 9.9 mm); pupil horizontally oval; tympanum (TYD 1.6 mm) distinct, rounded, 42 % of eye diameter, and equal to the disc of finger III (1.6 mm); tympanum-eye distance (TYE 0.9 mm), 56 % of tympanum diameter; supratympanic fold indistinct, extending from behind of eye to above the shoulder; parotoid glands absent; cephalic ridges absent; co-ossified skin on the head absent; vomerine teeth absent; tongue pear-shaped with deep notch posteriorly. Forelimbs short and gracile (Fig. 2; Fig. 3 A); forearm (FLL 5.9 mm) shorter than hand (HAL 8.9 mm), hand 67 % as long as foot (FOL 13.2 mm); forearm and hand length (14.8 mm) slightly smaller than half of SVL; dermal fringe along outer side of forearm absent; fingers I short and thin; fingers II, III (TFL 5.4 mm) and IV long and thin; relative length of fingers: I Coloration in life. Dorsal parts of head and dorsum, flank, loreal region, tympanic region and tympanum yellowish-olive, a dark-green irregular patch running from between eyes to shoulder, divaricating to the posterior part of the dorsum; upper and lower lip yellowish-olive with some creamy white spots; dorsal part of forelimb, thigh, tibia and foot yellowish-olive with transverse olive bands, forelimbs with 3 and hindlimbs with 6; lower part of flanks and tympanum with some dark brown spots; throat and margin of throat, chest, and belly marbled with white, ventral part of forelimbs and hindlimbs greyish-white; iris olive with brown reticulations; pupil periphery lined with yellow (Fig. 2 A, B, C). Coloration in formalin. Dorsal parts of head and dorsum, and upper part of flanks brownish grey; the dorsum with irregularly dark brown mark, a triangular spot between eyes bifurcating into two bands continuing posteriorly; lower flank beige with brown small spots; snout, canthus and loreal region, tympanum region, tympanum, and upper lip brown or brownish with white spots; vent region brown or brownish with white spots; forelimb, dorsal surface of thigh, tibia and foot light brown or grey with some dark brown bands; anterior and posterior part of thigh, and underneath the vent grey with small white spots; throat, margin of throat, chest and belly creamy white with brown marbling; ventral part of forelimbs and thighs creamy white; webbing greyish (Fig. 2 D, E, F). Male secondary sexual characters. Nuptial pad absent; internal subgular vocal sac present; lineae masculinae absent. Measurements of holotype. SVL, 30.0; HL, 12.0; HW, 11.2; MN, 10.0; MFE, 7.8; MBE, 3.4; IFE, 5.9; IBE, 9.9; IN, 2.9; EN, 3.0; EL, 3.8; NS, 1.6; SL, 4.7; TYD, 1.6; TYE, 0.9; IUE, 3.7; UEW, 2.8; HAL, 8.9; FLL, 5.9; TEL, 5.4; fd 3, 1.6; FL, 15.8; TL, 16.5; FOL, 13.2; FTL, 7.1; TFOL, 21.7; IMT, 1.3; ITL, 2.0; MTTF, 8.0; TFTF, 5.9; MTFF, 8.4; FFTF, 5.4. Variation. The tympanum almost equal to the width of the disc of finger III in males and obviously bigger than the disc of finger III in two female specimens (NHMG 20091002 and NHMG 201005008); morphometrics and coloration in the five paratypes are similar to those of the holotype. Measurements of type series are shown in Table 2. Habitat and distribution. The new species was found at elevations between 200��� 500 m. Between May and October vocalizing males were encountered at night (18:00��� 23:00 h, air temperature Etymology. This species is named after the locality in which it was collected. The suggested English name is the Nonggang small treefrog. Sequence divergence. Uncorrected sequence divergences between G. nonggangensis sp. nov. sequences and all homologous sequences available on the genus Gracixalus were greater than 6 %. The genetic distance is smallest between G. nonggangensis sp. nov. and G. jinxiuensis (EF 564525), at 6.1 %, whereas the intraspecific variation among G. nonggangensis sp. nov. range from 0 to 2.6 %. Among them, three specimens (NHMG 20091009 ��� 10, NHMG 111024) came from Nonggang station of Nonggang National Nature Reserve, which shared the same haplotype, whereas the genetically most divergent specimen (NHMG 1005046) came from Longrui station of Nonggang National Nature Reserve, at a distance of about 20 km from Nonggang station. Preliminary hypothesis of phylogenetic relationship. Based upon our preliminary molecular data, the genus Gracixalus formed a monophyletic group with well-supported values (BP= 98; BBP=1.00), and was divided two clades (Clade I and Clade II) with a strong support values (Fig. 4), consistent with Rowley et al. (2011). Clade I consists of G. gracilipes, G. supercornutus, G. quangi, and G. quyeti (= Clade I in Rowley et al. (2011)); Clade II consists of G. nonggangensis sp. nov., G. jinxiuensis, G. cf. jinxiuensis, Kurixalus cf. ananjevae, K. carinensis, K. odontotarsus and Gracixalus sp. (= Clade II in Rowley et al. (2011)). Among them, G. j i n x i u e n s i s did not form monophyletic groups, and indicated G. jinxiuensis group was likely a species complex. Based on the 16 S ribosomal RNA mitochondrial gene, we suggested that G. nonggangensis sp. nov. is sister to this group (G. cf. jinxiuensis, K. cf. ananjevae, K. carinensis, K. odontotarsus and Gracixalus sp.) with moderate supported values (BP= 90; BBP= 0.89). Comparisons. We compared G. nonggangensis sp. nov. with all rhacophorid treefrogs with a SVL under 45 mm from China and adjoining countries. Firstly, G. nonggangensis sp. nov. can be distinguished from all other species of the genus Gracixalus. Morphologically, the new species is most similar to G. quyeti (Nguyen et al. 2008). However, the species differs from G. quyeti by having a smooth, yellowish-olive dorsum, lacking tubercles (versus dorsal surface of head, back and upper portion of flanks covered with small sharp tubercles in G. quyeti) (Nguyen et al. 2008). Molecularly, phylogenetic trees indicated that G. nonggangensis sp. nov. did not cluster with G. quyeti in the same clade (Fig. 4), and the genetic distance (uncorrected p distance) between them reaches 12.7 %. G. nonggangensis sp. nov. differs from G. j i n x i u e n s i s, from Dayaoshan National Nature Reserve, Guangxi, by having a larger size in adults (SVL 29.9���35.3 mm male versus 23.5 mm male in G. jinxiuensis), and by having a yellowish-olive dorsum (versus brown dorsum in G. j i n x i u e n s i s) (Hu et al. 1978; Fei et al. 2009). G. nonggangensis sp. nov. differs from G. gracilipes by having a larger size in adults (SVL 29.9���35.3 mm male versus SVLG. gracilipes) and lacking white patch under the eye to the tympanum (present in G. gracilipes) (Delorme et al. 2005; Fei et al. 2009). G. nonggangensis sp. nov. can be distinguished from G. medogensis by having tibiotarsal articulation reaching to the tip of snout (versus to the front of eye in G. medogensis) (Fei et al. 2009). Furthermore, G. medogensis has a brown inverse ���V��� mark on the dorsum, and an orange mark on the flank of thigh (absent in G. nonggangensis sp. nov.) (Fei et al. 2009). G. nonggangensis sp. nov. differs from G. quangi by having a rounded snout (versus a triangularly pointed snout in G. quangi) and lacking black spots on the flanks and ventral surface of the thighs and a tibiotarsal projection (present in G. quangi), and having a larger size in adults (SVL 29.9���35.3 mm male versus SVLG. quangi) (Rowley et al. 2011). G. nonggangensis sp. nov. differs from G. supercornutus by having a larger size in adults (SVL 29.9���35.3 mm male versus SVLG. supercornutus) and lacking prominent tubercles or spines on the whole dorsum and the upper eyelids (versus present in G. supercornutus) (Orlov et al. 2004). The yellow-olive dorsum, finger webbing absent, toes one-third webbed, and lacking vomerine teeth and macroglands separate G. nonggangensis sp. nov. from all species of Buergeria, Kurixalus, Polypedates and Rhacophorus. Buergeria japonica has a brown dorsum (Fei et al. 2009). Buergeria oxycephala has fingers lightly webbed and toes fully webbed (Fei et al. 2009). The species of Chiromantis have an opposable finger (versus noopposable fingers in G. nonggangensis sp. nov.). Feihyla palpebralis has a distinct, broad silvery white bands below the eyes, and a nuptial pad on finger I and II. Feihyla fuhua has a long and wide white streak from below eye to the base of shoulder (Fei et al. 2009; Fei et al. 2010). Kurixalus carinensis has distinct supratympanic fold and macrogland. Kurixalus ordontotarsus has dermal fringes and tubercles on the limbs, and sharp tubercles on the dorsum (Ye et al. 1993; Nguyen et al. 2008). Kurixalus eiffingeri has some white glands in the forearms and hindlimbs, and well developed vomerine teeth (Fei et al. 2009). Kurixalus bisacculus is dark brown with only a tubercle at the heel (Taylor 1962). Kurixalus verrucosus have a brown dorsum with only a tubercle at the heel (Boulenger 1893). Liuixalus hainanus has brown oval mark on the dorsum (Liu & Wu 2004). Liuixalus ocellatus has a brown dorsum with some glands (Liu & Hu 1973). Liuixalus romeri has a small size (SVL 15���18 mm in male versus 29.9���35.3 mm in male for G. nonggangensis sp. nov.) (Smith 1953). Philautus abditus, Philautus cardamonus and Philautus kempii have a brown dorsum (Inger et al. 1999, Ohler et al. 2002, Annandale 1912). Philautus maosonensis has a brown dorsum, and truncated snout (Bourret 1937). Philautus truongsonensis has a brownish dorsum, an obtusely pointed snout (Orlov & Ho 2005). Philautus tytthus is dark greyish (Smith 1940). The species of Polypedates have vomerine teeth, and finger webbing. Rhacophorus arvalis has a large size (SVL ranging from 42���44 mm in male versus 29.9���35.3 mm in male for G. nonggangensis sp. nov.), and finger webbing. Rhacophorus chenfui has white flank lines that clearly divide dorsal and ventral portions and purple flanks in life (Fei et al. 2009). Rhacophorus dugritei and Rhacophorus hungfuensis have vomerine teeth and finger webbing. Rhacophorus laoshan lack a green dorsum and has vomerine teeth. Rhacophorus leucofasciatus and Rhacophorus minimus have vomerine teeth, and a white line running from snout to the lateral of body and ending before the hindlimbs. Rhacophorus moltrechti has finger webbing. Rhacophorus rhodopus has vomerine teeth, and finger and toe fully webbed (Rao et al. 2006; Fei et al. 2009). Rhacophorus taipeianus has green dorsum with small glands, and finger wedding. Rhacophorus translineatus has triangularly pointed snout, and finger webbing. Rhacophorus verrucopus has a tibiotarsal projection, and finger webbing. Rhacophorus yaoshanensis has weak tuberculous dermal ridge along outer sides of forearm and tarsus, and finger webbing (Fei et al. 2009). Rhacophorus yinggelingensis has a lager size (SVL> 40 mm in male versus 29.9���35.3 mm in male for G. nonggangensis sp. nov.), and the front of thigh is yellow and red-tinged, and the rear of thigh and inner side of tibia are red (Chou et al. 2007). Raorchestes kakachi has indistinct tympanum and blotches of dark brown on flanks (Seshadri et al. 2012). Raorchestes longchuanensis and Raorchestes menglaensis have a brown dorsum (Yang & Li 1978; Kou 1990). Raorchestes parvulus has a brownish grey dorsum (Boulenger 1893). Raorchestes resplendens has a bright orange to reddish coloration, multiple macroglands on the body and extremely short limbs (Biju et al. 2010). Theloderma andersoni, Theloderma asperum, Theloderma baibengensis, Theloderma kwangsiensis, Theloderma moloch, Theloderma lateriticum, Theloderma rhododiscus, and Theloderma stellatum lack a green dorsum or have some glands on the dorsum (Ahl 1927; Boulenger 1886; Jiang et al. 2009; Bain et al. 2009; Liu & Hu 1962; Taylor 1962; Fei et al. 2009)., Published as part of Mo, Yunming, Zhang, Wei, Luo, Yu, Zhou, Shichu & Chen, Weicai, 2013, A new species of the genus Gracixalus (Amphibia: Anura: Rhacophoridae) from Southern Guangxi, China, pp. 61-72 in Zootaxa 3616 (1) on pages 64-69, DOI: 10.11646/zootaxa.3616.1.5, http://zenodo.org/record/283369