Amolops gudao Yu, Wu, Lu & Che 2025, sp. nov.
Creators: name: Wu, Yun-He
name: Yu, Zhong-Bin
name: Kilunda, Felista Kasyoka
name: Lu, Chen-Qi
name: Li, Jia-Hua
name: Li, Yun-Peng
name: Shi, Yi-Juan
name: Che, Jing
Publisher: name: Zenodo
Year: 2025
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Ranidae
Amolops
Amolops gudao

description: Amolops gudao Yu, Wu, Lu & Che sp. nov.
Figs 1, 2, 3, 4, 5
Type material examined.
Holotype • Adult male (KIZ 053662), from Baihualing Village, Longyang, Baoshan, Yunnan Province, China (25.301208 ° N, 98.788815 ° E; elevation 1793 m a. s. l.), collected by Zhong-Bin Yu, Dong An, Tian-En Chen on 19, July, 2021. Paratypes • One adult male (KIZ 053663), two adult females (KIZ 053664 and KIZ 053665); collected at the same locality and with the same collection information as the holotype.
Diagnosis.
The new species is recognized as a member of the A. viridimaculatus group based on molecular phylogenetic analyses and can be distinguished from its groups by a combination of the following characters (Suppl. material 3): (1) medium body size (SVL 56.7–58.6 mm in males and 63.6–64.8 mm in females); (2) vomerine teeth developed, on two short oblique between choanae, “ 八 ” - shaped, almost equal in distance from each other as to choanae; (3) supratympanic fold indistinct; (4) true dorsolateral folds absent, discontinuous glandular dorsolateral fold from rear of eye to near vent present; (5) tongue cordiform, 1 / 2 notched posteriorly; (6) maxillary teeth developed; (7) circummarginal grooves present on tips of outer three fingers, absent on first finger; (8) inner metacarpal tubercle rounded, outer metacarpal tubercle indistinct; (9) tibiotarsal articulation of adpressed limb reaching the anterior corner of eye; (10) a black stripe below edge of the canthus rostralis extending from the nostril across the eyes, along dorsolateral glandular folds to near vent; (11) rictal gland absent; (12) iris distinctly bicolored, green in upper 1 / 4 and reddish brown in lower 3 / 4, black reticulations throughout; (13) vocal sac absent in males; (14) male with orange nuptial pad at the base of first finger.
Description of holotype.
(measurements in Table 1) KIZ 053662, a medium-sized adult male (SVL 58.6 mm); body relatively robust; head slightly longer than wide (HL 18.2 mm; HW 17.7 mm); outline of the snout rounded in dorsal, ventral and lateral views, projecting beyond lower jaw, its length longer than diameter of eye (SL / ED 131.7 %); loreal region concave and oblique; canthus rostralis distinct; interorbital space flat, shorter (IOS 4.9 mm) than upper eyelid width (UEW 5.2 mm) and internarial distance (INS 6.5 mm); nostril oval, laterally orientated, closer to anterior corner of eye (NED 3.8 mm) than tip of snout (SND 4.1 mm); tympanum indistinct, circular in shape, relatively small (TD / HL 13.7 %), tympanic rim slightly elevated, indistinct; eyes relatively large (ED / HL 34.6 %), slightly protuberant in dorsal view (Fig. 3 A) and notably protruding in profile (Fig. 3 B), eye diameter longer than snout length (SL / ED 131.7 %); pupil horizontal (Fig. 3 B); vomerine teeth developed, on two short oblique rows between choanae, “ \ / ” - shaped, almost equal in distance from each other as to choanae; tongue attached anteriorly, cordiform, 1 / 2 notched posteriorly; choanae oval; maxillary teeth developed; indistinct tooth-like projection on anteromedial edge of mandible.
Forelimbs moderately long and robust, forearms significantly enlarged relative to upper arms; forelimb and hand length (LAHL 30.6 mm) longer than 1 / 2 body size (LAHL / SVL 52.2 %); relative length of fingers: FI <FII <FIV <FIII; tips of all four fingers expended into discs, disc on finger III approximately equal to 1 / 2 snout length (FD 3 4.2 mm; FD 3 / SL 50.6 %); circummarginal grooves present on tips of outer three fingers, absent on first finger; subarticular tubercles prominent and oval, formula 1, 1, 2, 2; supernumerary tubercles present at the base of each finger; webbing between fingers absent; narrow lateral fringes of fingers III and IV; inner metacarpal tubercle rounded, outer metacarpal tubercle indistinct (Fig. 3 D).
Hindlimbs long and robust, almost three times longer than the forearms and nearly two times longer than body length (LAHL / HLL 31.0 %; SVL / HLL 59.4 %); femoral length shorter than the tibia length (FEM / TIB 95.0 %) and the foot length (FEM / FTL 88.8 %); tibiotarsal articulation of adpressed limb reaching the anterior corner of eye when hindlimb is stretched alongside the body; the heels slightly overlapping when the tibias are perpendicular to the body axis; relative toe lengths: TI <TII <TIII <TV <TIV; toes with narrow lateral fringes, toe IV and postaxial side of toe V relatively wider; tips of all toes expanded into discs (TD 1 2.9 mm; TD 2 3.4 mm; TD 3 3.2 mm; TD 4 2.2 mm; TD 5 1.7 mm), toe discs relatively smaller than those of fingers; toe discs with circummarginal grooves; toes fully webbed except for fourth toe, in which web reaches beyond distal subarticular tubercle; subarticular tubercles oval and distinct, formula 1, 1, 2, 3, 2; supernumerary tubercles absent; inner metatarsal tubercle elongate (IMT / TI 68.9 %), outer metatarsal tubercle absent (Fig. 3 E).
Skin on dorsal surface of head, body and limbs smooth; flank of body, ventral surface of head and limbs, throat, chest and abdomen relatively smooth; temporal region without any tubercles; supratympanic fold indistinct; true dorsolateral folds absent, but discontinuous series of glands along dorsolateral junction of body (dorsolateral glandular folds) present, extending from rear of eye to near vent; posterior angle of the jaw with dense tubercles; rictal gland absent (Fig. 3).
Color of holotype in life.
The ground coloration of dorsum vivid green, interspersed with irregular black spots; ventral surface of the head, throat, chest and belly mostly yellow-green, posterior abdomen with dense yellow dots; ventral surface of thighs, tibia, and tarsus brown, scattered with green spots and blotches; lateral surfaces of the body vivid green; a black stripe below edge of the canthus rostralis extending from the nostril across the eyes, along dorsolateral glandular folds to near vent; the dorsal surfaces of limbs with irregular black transverse bands, mottled with yellow-green dots, and the bands much more distinct on the hindlimbs, bands relatively incomplete on the forelimbs; iris distinctly bicolored, green in upper 1 / 4 and reddish brown in lower 3 / 4, black reticulations throughout; foot webbing brown; ventral surface of outer three fingers discs and outer three toes discs, metatarsal tubercle, subarticular tubercles of toes gray; ventral surface of first finger disc, inner two toes discs, subarticular tubercles and supernumerary tubercles of fingers orange yellow (Fig. 3).
Color of holotype in preservative.
After one year of storage in ethanol, dorsum metallic blue, with small scattered blackish spots; transverse cross-bars on dorsal surfaces of hands, shanks, tarsus, and feet still clear and turning black; a black stripe below edge of the canthus rostralis extending from the nostril across the eyes, along dorsolateral glandular folds to near vent still clear; chest, throat and ventral surface of head fading to dark gray; ventral surface of limbs fading to cream-yellow, scattered with brown pigmentations; abdomen brown, gray around abdomen; posterior angle of the jaw and posterior surface of thigh around vent with dense tubercles more distinct; digit tips, nuptial pads, supernumerary tubercles, subarticular tubercles, and metatarsal tubercle fading to cream-yellow or grayish-white; foot webbing cream-yellow (Fig. 4).
Secondary sexual characters.
Adult female specimens have larger body sizes than adult male specimens (SVL 63.6–64.8 mm vs 56.7–58.6 mm, Table 1). Adult males possess orange nuptial pads covering the dorsal surface of base of the first finger; absence of vocal sacs in males. Forearms of male are slightly enlarged relative to the upper arms (Fig. 3).
Morphological variation.
The overall morphology of paratypes agrees with the holotype description with the following exceptions. KIZ 053663 (male): The ground coloration of dorsum vivid green, with larger and more irregular black markings mottled on the dorsal area, scattered with a few yellow spots; complete transverse cross-bars on dorsal surfaces of limbs, cross-bands with distinct yellow edge. KIZ 053664 (female): compared to holotype and paratypes, larger irregular black markings mottled on dorsal area, scattered with a few yellow spots; complete transverse cross-bars on dorsal surfaces of hindlimbs, cross-bands without distinct yellow edge; flanks present two black specks. KIZ 053665 (female): compared to other holotype and paratypes, smaller irregular black markings mottled on dorsal area, connected to form reticulations throughout; upper lips with three dark vertical bars; flanks mottled with seven black specks; dorsal surfaces of hindlimbs without complete transverse cross-bars, but scattered with some black specks; complete transverse cross-bars on dorsal surfaces of hindlimbs (Fig. 5).
Distribution.
Currently, this species is only known from Baihualing village, Longyang, Yunnan, China.
Ecology and habitat.
This nocturnal species inhabits wide and swift mountain streams. There were many large stones covered with moss in the stream. The frog was observed squatting on big rocks in the streams or on banks during our survey from 21: 00 to 24: 00 in July. When disturbed, it immediately jumps into the water. The vegetation on both sides of the stream was dense, surrounded by broad-leaved forests and bamboo forests. The adults were only found on July 19. The adult males displayed orange nuptial pads and the females were gravid during specimen collection. The tadpoles of this species were found on July 8 at the collection site. Therefore, we speculate that the breeding season of this species occurs around July-August. This species is sympatric with A. viridimaculatus (Jiang, 1983) and its congeners. The new species was also found in sympatry with Leptobrachella alpina (Fei, Ye & Li, 1990), Oreolalax jingdongensis (Ma, Yang & Li, 1983), Bufo tuberospinius (Yang, Liu & Rao, 1996), Boulenophrys lushuiensis (Shi, Li, Zhu, Jiang, Jiang & Wang, 2021), Leptobrachium huashen Fei & Ye, 2005, Nanorana yunnanensis (Anderson, 1879), and Zhangixalus burmanus (Andersson, 1939).
Comparisons.
The new species was compared with the other species of the A. viridimaculatus species group (Suppl. material 3; Andersson 1938; Biju et al. 2010; Nidup et al. 2016; Qi et al. 2019; Che et al. 2020; Mahony et al. 2022; Saikia et al. 2022; Hou et al. 2023; Wu et al. 2024 a).
Amolops gudao sp. nov. is significantly different from A. wangyali by smaller body size, male SVL 56.7–58.6 mm and female SVL 63.6–64.8 mm (vs SVL 71.4–76.7 mm in males and 80.5–89.6 mm in females), rictal gland absent (vs a distinct patch of rictal glands at rear of jaw on either side), iris distinctly bicolored, green in upper 1 / 4 and reddish brown in lower 3 / 4, black reticulations throughout (vs pupil with near continuous pale metallic green border, remaining iris mottled metallic green and chocolate brown, more green than brown on dorsal 1 / 3 and ventral most portions of iris), male with orange nuptial pad at the base of first finger (vs nuptial pads yellowish-gray dorsally, dark gray ventrally); from A. chanakya by smaller adult male size, male SVL 56.7–58.6 mm (vs 76.4 mm), supratympanic fold indistinct (vs supratympanic fold strong), the ground coloration of dorsum vivid green, interspersed with irregular black spots (vs dorsal color dull brick-red, spotted with irregular cocoa-brown spots, these cocoa-brown spots enclosing a number of smaller dull brick-red spots), absence of vocal sacs in males (vs vocal sac externally visible); from A. tawang by smaller adult male size, male SVL 56.7–58.6 mm (vs 82.5 mm), tibiotarsal articulation of adpressed limb reaching the anterior corner of eye (vs tibio-tarsal articulation reaches to snout), supratympanic fold indistinct (vs supratympanic fold strong), the ground coloration of dorsum vivid green, interspersed with irregular black spots (vs dorsal color olive-green, spotted with large, irregular shaped dark-brown spots, brown spots enclosing a number of small olive-green dots), absence of vocal sacs in males (vs vocal sac faintly visible); from A. kaulbacki by smaller body size, male SVL 56.7–58.6 mm and female SVL 63.6–64.8 mm (vs SVL 72.6–82.6 mm in males and 82.7–87.2 mm in females), tibiotarsal articulation of adpressed limb reaching the anterior corner of eye (vs tibio-tarsal articulation reaches to the tip of snout), iris distinctly bicolored, green in upper 1 / 4 and reddish brown in lower 3 / 4 (vs iris vivid green with irregular dark patterns), discontinuous glandular dorsolateral fold (vs absent); from A. viridimaculatus by smaller body size, male SVL 56.7–58.6 mm and female SVL 63.6–64.8 mm (vs 72.7–82.3 in males and 83.0– 94.3 in females), discontinuous glandular dorsolateral fold (vs absent), the ground coloration of dorsum vivid green, interspersed with irregular black spots (vs dorsum and flank with nearly round green or yellowish green spots, scattered with small green spots); from A. beibengensis by smaller body size, male SVL 56.7–58.6 mm and female SVL 63.6–64.8 mm (vs SVL 75.8 mm male and 90.2–93.2 mm in females), discontinuous glandular dorsolateral fold (vs absent), male with orange nuptial pad at the base of first finger (vs white nuptial pads), supratympanic fold indistinct (vs distinct, wide and thick); from A. formosus by smaller adult female size, SVL 63.6–64.8 mm (vs SVL 79.4–83.7 mm), glandular dorsolateral fold (vs absent), male with orange nuptial pad at the base of first finger (vs developed white nuptial pads); from A. himalayanus by smaller body size, male SVL 56.7–58.6 mm and female SVL 63.6–64.8 mm (vs SVL 72.8–76.7 mm males and 80.5–89.0 mm in females), vocal sac absent in males (vs externally visible vocal sacs present), the ground coloration of dorsum vivid green, interspersed with irregular black spots (vs dark brown, interspersed with irregular yellow spots); from A. pallasitatus by smaller adult female size, female SVL 63.6–64.8 mm (vs 70.6–72.3 mm in females), glandular dorsolateral fold (vs absent), the ground coloration of dorsum vivid green, interspersed with irregular black spots (vs dorsum yellow-green, with irregular dark brown blotches without margins), rictal gland absent (vs rictal gland prominent and ellipsoidal); from A. yangi by larger body size, male SVL 56.7–58.6 mm and female SVL 63.6–64.8 mm (vs SVL 46.3–51.8 mm males and 51.5 mm in female), three metacarpal tubercles, inner metacarpal tubercle long, outer metacarpal tubercle relatively small, oval, median one rounded (vs inner metacarpal tubercle rounded, outer metacarpal tubercle indistinct), a black stripe below edge of the canthus rostralis extending from the nostril across the eyes, along dorsolateral glandular folds to near vent (vs a black stripe below edge of the canthus rostralis extending from the snout tip across the eyes, to the anterior edge of supratympanic fold), toes with narrow lateral fringes, toe IV and postaxial side of toe V relatively wider (vs narrow lateral fringes of preaxial side of toe I and postaxial side of toe V); from A. longimanus by larger body size, male SVL 56.7–58.6 mm and female SVL 63.6–64.8 mm (vs SVL 30 mm), eye diameter (ED / SL 70.9 % – 75.9 %) shorter than snout length (vs snout approximately as long as the eye diameter, ED / SL 97.7 %), nostrils closer to anterior corner of eye than to tip of snout (vs nostrils a little nearer to tip of snout than the eye), hand length ~ 2 / 3 of forelimb (vs the hand nearly 1 / 2 length of the whole forelimb); from A. medogensis by smaller body size, male SVL 56.7–58.6 mm and female SVL 63.6–64.8 mm (vs SVL 95.0 mm males and 72.4–96.9 mm in females), supratympanic fold indistinct (vs distinct, wide and thick), tibiotarsal articulation of adpressed limb reaching the anterior corner of eye (vs beyond tip of snout), circummarginal grooves present on discs of outer three fingers, absent on first finger (vs discs of all fingers with circummarginal grooves); from A. wangyufani by smaller body size, male SVL 56.7–58.6 mm and female SVL 63.6–64.8 mm (vs SVL 68.3–69.0 mm males and 83.4 mm in female), discontinuous glandular dorsolateral fold (vs absent), tibiotarsal articulation of adpressed limb reaching the anterior corner of eye (vs beyond tip of snout), the ground coloration of dorsum vivid green, interspersed with irregular black spots (vs brown dorsal color, head and body with green markings), male with orange nuptial pad at the base of first finger (vs pale gray nuptial pads); from A. nidorbellus by smaller body size, male SVL 56.7–58.6 mm and female SVL 63.6–64.8 mm (vs SVL 76.4–82.3 mm males and 85.4–98.0 mm in females), the ground coloration of dorsum vivid green, interspersed with irregular black spots (vs dorsally brown with small irregularly arranged cobalt green spots), tympanum indistinct (vs distinct).
Etymology
Baihualing, where the new species occurs, lies on the historical passage of the famous Southern Silk Road. The specific epithet gudao is derived from the Chinese alternative name for the Southern Silk Road. We propose the English common name Gudao Cascade Frog and the Chinese common name Gǔ Dào Tuān Wā (古道湍蛙).
description: Published as part of Wu, Yun-He, Yu, Zhong-Bin, Kilunda, Felista Kasyoka, Lu, Chen-Qi, Li, Jia-Hua, Li, Yun-Peng, Shi, Yi-Juan & Che, Jing, 2025, A new species of the genus Amolops (Anura, Ranidae) from the Gaoligong Mountains, China, pp. 349-367 in ZooKeys 1227 on pages 349-367, DOI: 10.3897/zookeys.1227.131357