Description of a new toad of Megophrys Kuhl & Van Hasselt, 1822 (Amphibia Anura: Megophryidae) from western Yunnan Province, China Shi, Shengchao Li, Donghui Zhu, Wenbo Jiang, Wen Jiang, Jianping Wang, Bin Zootaxa 2021 2021-03-15 4942 3 351 381 Shi & Li & Zhu & Jiang & Jiang & Wang, 2021 Shi & Li & Zhu & Jiang & Jiang & Wang 2021 [151,431,1481,1508] Amphibia Megophryidae Megophrys Animalia Anura 9 360 Chordata species lushuiensis sp. nov.   Figs. 3–6; Tables 1, 3–5     Holotype. Adult male, CIBYN201909290, collected from Gutan River in Luzhang Town, Lushui County, Nujiang Prefecture, Yunnan Province, China( 25.977066° N, 98.795983° E, ca.  2358 ma.s.l.) by SC Shi, WB Zhu, and L Fuon  27 thSeptember 2019.    Paratypes. One adult male( CIBYN201909283), collected from Bashan, Luzhang Town, Lushui County( 25.953588° N, 98.734431° E,  1986 m), other two toadlets ( CIBYN201909288–289) collected from Langbazhai Village, Luzhang Town( 25.95118° N, 98.768903° E,  2006 m), by SC Shi, WB Zhu, and LFuon  27 thSeptember, 2019.   Other specimens checked. Fivetadpoles. CIBYN201909119, collected from Haobadi, Laowo Town, Lushui County( 25.808908° N, 99.107779° E,  2363 m) on  23 rdSeptember, 2019.  Twotadpoles CIBYN201909171–172, collected from Bashan, Luzhang Town, Lushui Countyon  27 thSeptember 2019.  CIBYN201909175, collected from the same site together with the holotype.  CIBYN201909176, collected from a small stream near the Gutan River of Luzhang Town, Lushui County( 25.958701° N, 98.717995° E,  2399 m) on  27 thSeptember 2019.  Alltadpoles collected by SC Shi, WB Zhu, and L Fu.   Etymology.The species name  lushuiensisis inferred from the typelocality Lushui County, Yunnan Province, China.   Suggested vernacular name.Lushui horned toad (English), and Lu Shui Jiao Chan (aeḳfflffi, Chinese).   Diagnoses.  Megophrys lushuiensis  sp. nov.is assigned to the genus  Megophrys sensu latobased on molecular phylogenetic analyses and the following morphological characters: canthus rostralis well-developed; a small horn’- like tubercle at edge of upper eyelid present; supratympanic fold distinct; axillary glands small, on sides of the breast; oral disc of tadpoles funnel-like; mouth of tadpoles lacking transverse rows of teeth; head length more than 25% of body size; upper jaw protruding beyond the margin of the lower jaw; no skin fold on back of head; maxillary teeth present; tympanum distinct; hind legs long and thin.   Megophrys lushuiensis  sp. nov.is identified from its congeners by a combination of the following characters: moderate body size (SVL 31.0– 34.8 mmin males, n = 2); vomerine ridge weak, vomerine teeth absent; dorsal skin relatively smooth; tongue slightly notched behind; tympanum rounded and relatively large, 0.54 times of eye length; horn’-like tubercle on edge of upper eyelid small; tibio-tarsal articulation reaches middle eye, tibia relatively long, ca. 0.56 times of body length; finger tips rounded, not expanded to small pad; toes with narrow fringes and rudimentary webbing; ventral hindlimbs semitransparent purplish with greyish white pigments; ventral body scattered with distinct dark patches in the middle.    Holotypedescription. Figs. 3and 4; Table 2. Measurements in mm. Body size small (SVL 34.8) and relatively robust; protruding beyond cloaca small, not visible from ventral view. Head moderate, longer than wide (HW 12.5, HL 12.8, IFE 6.2, IBE 9.6); snout rounded in dorsal view, slight projecting in profile, protruding beyond lower jaw; rostral appendage absent (SL 4.3); canthus rostralis distinct (right side wounded); loreal region concave; nostril oval, nearly oval, closer to eye than snout (SN 2.3, EN 2.1), distance between nostrils slightly larger than distance between upper eyelids (IN 4.1, IUE 3.9); tympanum slightly larger than half of eye diameter (EL 3.9, TYD 2.0), and larger than eye-tympanum distance (TYE 1.6); tympanum rounded, slightly raised above surrounding, upper edge conceal with supratympanic ridge; interorbital space flat, wider than upper eyelid (UEW 3.4); pineal ocellus not observed; vomerine ridges weak, vomerine teeth absent; maxillary teeth present; tongue weakly notched behind, medial lingual process absent. Forelimbs. Forearm moderately long, slightly enlarged than upper arms, shorter than hand (FAL 6.7, HAL 7.6); fingers slender, without webbing and lateral fringes, subarticular tubercles present on finger I and II, absent on other fingers, base of first finger distinct larger than the second; inner and outer metacarpal tubercles slightly raised above hand, separated from each other; finger length formula II<I<IV<III; tip of fingers rounded, slightly larger than last phalanges, not expand to small pad (FIIIW 0.6, FIVW 0.6). Hindlimbs thin and long, tibio-tarsal articulation reaches middle eye when leg stretched forward; thighs slightly shorter than shanks but longer than feet (TL 16.6, SHL 17.0, FOL 13.1, TFOL 21.1); toes long, relatively lengths I <II <V <III <IV, rudimentary webbed, webbing formula: I 2–2 ¾ II 2–3¾ III 3–4¼IV 4–3V, with narrow lateral fringes, tips rounded and not expanded; dermal ridges continuously present on ventral surface of toes; subarticular tubercles absent; outer metatarsal tubercle absent, inner metatarsal tubercle oval, small, positioned by the base of first toes (IMT 1.5). Skin. Dorsal surface of head, body and limbs relatively smooth, scattered with small granular tubercles all over; tip of most granules with one keratinized spinule. Lateral head scattered with several granules below supratympanic ridges and around tympanum. A small tubercle present on outer edges of upper eyelids. Supratympanic ridges thin before and above departure with tympanum, expand into a drop–shaped of water after, and curve above tympanum. Two longitudinal skin folds present on dorsolateral body, extending from above posterior supratympanic ridges to middle body. A longitudinal “>-<” shaped skin ridge present on middle dorsum. Skin ridge on head between eyes absent. Four short near transverse skin ridges present on dorsal thighs and shanks. Several larger flat tubercles present on flanks with width of two or three finger tips. Ventral surface of body and limbs smooth. Pectoral glands flat, small, with the size of first fingertip. Femoral glands small, closer to cloaca than outer edge of knee. Spinous small granules present around cloaca. Coloration in life. Fig. 3. Dorsal surface basically saddle brown. A dark brown triangular pattern presents between eyes, edged with lighter color, posterior part of triangular pattern on head fused with blur “>-<” shaped dark brown pattern on dorsum. Dorsal limbs with several wide blur transverse brown bars. Lateral head brown; upper edge of supratympanic ridges light brown, while lower edge dark brown; tympanum and skin between tympanum to eye dark brown; a wide dark brown bar present under eyes on upper lips. Lateral body tan; tubercles on flanks lighter colored; groins dark orange. Ventral body lighter than dorsal. Gular and chest purplish grey, the longitudinal stripes extend from mouth corner to ventral upper arm and the central longitudinal stripe on throat are ambiguous; two small white patches present on lower lips below snout. Abdomen ivory, four or five large dark patches present on both sides of ventrolateral body; about a dozen of distinct black irregular patches present on middle belly. Skin on lower arm semitransparent purplish with greyish white pigments, inner forearms with two large black patches; skin on ventral hindlimbs semitransparent purplish with greyish white pigments. Ventral hand dim grey; inner and outer palm tubercles and dark orange; ventral surface of fingers stained with little dark orange, more distinct on finger I. Ventral feet dim grey, stained with dark orange. A black patch present around cloaca, not seem from ventral view. Pectoral and femoral glands cream white. Iris copper above, light grey below.   FIGURE 3.Photos of the holotype CIBYN201909290 of  Megophrys lushuiensis  sp. nov.in life. A, dorsal view. B, ventral view. C, lateral view. D, ventral view of hand. E, ventral view of foot. F, maxillary view, noting vomerine ridges weak and vomerine teeth absent. Coloration in preserve. Fig. 4. Described when holotypefixed in formalin and then preserved in 75% ethanol for three months. Coloration lighter than in life, dorsal surface mostly grey, dark brown patterns on head, body and hindlimbs dim grey. Black patches on body and limbs remained black. Dark orange color on groin and ventral hand remained. Ventral surface whiter than in life. Purplish color faded.  Secondary sexual characters.In breeding male, nuptial pads present on inner dorsal surface of the first two fingers, tiny spines on the nuptial pads dense; single subgular vocal sac present in male.  Variation. Fig. 5; Table 2. Adult male paratypeCIBYN201909283, with a smaller body size (SVL 31.0); deeper coloration; ventral hindlimbs covered with lesser greyish white pigments; ventrolateral side of belly with a large and long dark patch on both side; skin ridges on dorsolateral body connect with the “Λ” shaped skin ridge on middle dorsum. One of two toadlets collected with light brown dorsal coloration. Patterns on dorsum of two toadlets collected not distinguishable.   TABLE 2.Measurements (in mm) of two adult male and two juvenile specimens of  Megophrys lushuiensis sp. nov.    Voucher  CIBYN201909290  CIBYN201909283  CIBYN201909288  CIBYN201909289   Status Holotype Paratype Paratype Paratype   Sex Male Male Male Male   Development Adult Adult Juvenile Juvenile  SVL 34.8 31.0 17.8 18.7  HW 12.5 11.9 6.8 7.0  HL 12.8 12.4 7.0 6.6  SL 4.3 4.1 2.5 2.6  SN 2.3 2.3 1.2 1.5  EN 2.0 1.9 1.1 1.1  IN 4.1 3.9 2.5 2.6  EL 3.9 3.8 2.3 2.5  IUE 3.0 3.2 2.3 2.5  UEW 3.4 2.4 1.5 1.5  IFE 6.2 5.5 4.0 3.7  IBE 9.6 8.7 5.9 6.2  TYD 2.0 2.1 1.2 1.1  TYE 1.6 1.8 0.9 1.0  FAL 6.7 7.3 4.0 3.8  HAL 8.8 8.2 5.2 5.1  TL 16.6 16.2 9.0 9.2  SHL 17.0 17.5 9.9 10.1  TFOL 24.5 24.2 13.7 13.3  FOL 15.6 15.5 8.6 8.9  IMT 1.5 1.3 0.6 1.1  FIIIW 0.6 0.6 0.3 0.4  FIVW 0.6 0.5 0.3 0.3  HLL 54.4 55.2 30.8 30.7  Tadpoles. Fig. 6; Table 3. Measurements in mm. Description based on the tadpole CIBYN201909171 ( Fig. 5) at Gosner stage 32. Body elongated, length 2.8 times of width (BL 12.1, BH3.9, BW4.4); head as wide as trunk; eyes moderate (ED 1.1), rounded, positioned on dorsolateral head, widely separated (PP 3.4); nares small, much closer to pupils than tip pf snout (NE 0.7, RN 2.5), internasal distance much shorter than interpupil distance (IND 2.4); oral disc funnel like, anterodorsal positioned, width 1.7 times of body width (ODW 7.6); submarginal papillae on lips in shape of short bar near the opening of mouth and oval near fringes, major axis of submarginal papillae pointing to the opening of mouth, four or five rows on middle lower lip, 3 rows of oval submarginal papillae on both sides of upper lips; keratodonts absent; spiracle in right-handed helix from ventral view, the spiracular tube not protruding beyond body wall, positioned 58% of the distance between tip of the snout and trunk-tail junction (SS 7.1), and below the horizontal mid trunk line; tail long, length 73% of total length (TAL 30.7); dorsal fin slowly arise above trunk-tail junction, maximum height 35% of body height (UF 1.4); ventral fin connected to the trunk, height subequal to dorsal fin, 33% of maximum body height (LF 1.3); maximum tail muscle height 94% of maximum body height, maximum tail muscle width 61% of maximum body width (TMW 2.7, TMH 3.6); anal siphon opens medially.   FIGURE 4.Photos of the holotype specimen CIBYN201909290 of  Megophrys lushuiensis  sp. nov.A, dorsal view. B, ventral view. C, lateral view of head. D, ventral view of hand. E, ventral view of foot. When alive, dorsal body and tail brown, without distinct patterns. lateral body light brown, skin on lower half body semitransparent, deep greyish; lateral tail light brown with dozens of small dark brown patches on tail; fins semitransparent. Ventral body semitransparent, scattered with silver tiny patches, deep grey intestines visible. A golden ring present around pupil, golden pigments scattered around the ring. In preservation, coloration deeper; brown pigmentation on body more prominent than when alive; ventral body and tail fin semitransparent white with brown pigments; tiny silver patches on belly less distinct than when alive.  Comparisons.  Megophrys lushuiensis  sp. nov.is phylogenetically assigned to the subgenus  Panophrysand can be distinguished from its congeners of the subgenus by morphological characters as shown in Table 5. We also compared the new species with other species of  Megophrys. By tympanum distinct large,  Megophrys lushuiensis  sp. nov.differs from species of subgenus  Atympanophrys, i.e.,  M. gigantica,  M. nankiangensis,  M. shapingensis, and  M. wawuensis(vs. absent, concealed or very small in the latter). By maxillary teeth present,  Megophrys lushuiensis  sp. nov.differs from species of subgenus  Ophryophryne, i.e.,  M. elfina,  M. gerti,  M. hansi,  M. koui,  M. microstoma, and  M. synoria(vs. absent in the latter). By head not wide flat and hind limbs long,  Megophrys lushuiensis  sp. nov.differs from  M. carinense,  M. chuannanensis,  M. feae,  M. intermedia,  M. orientalis,  M. popei, and  M. platyparietus(vs. head wide flat and hind limbs short in the latter). By lacking a single, wide and flat palpebral projection on the edge of the upper eyelid,  Megophrys lushuiensis  sp. nov.differs from species of subgenus  Megophrys, i.e.,  M. lancip,  M. montana,  M. parallela, and most of species of subgenus  Pelobatrachus, i.e.,  M. baluensis,  M. edwardinae,  M. kobayashii,  M. ligayae,  M. nasuta, and  M. kalimantanensis(vs. present in the latter). By lacking rostral appendage,  Megophrys lushuiensis  sp. nov.differs from  M. stejnegeri(vs. having less rostral appendage in the latter). By having a small horn-like tubercle at edge of upper eyelid,  Megophrys lushuiensis  sp. nov.differs from  M. dringi(vs. very large in the latter). By body size moderate,  Megophrys lushuiensis  sp. nov.differs from  M. feii(two males SVL 31.0– 34.8 mmin the new species vs. four males SVL 24.5–25.1 inthe latter). By finger tips not expanded, the new species differs from  M. feii(vs. expanded in the latter).   FIGURE 5.Color variation in  Megophrys lushuiensis  sp. nov.in life. A and B, dorsolateral and ventral view of adult male paratype CIBYN201909283, SVL 31.0 mm, respectively. C and D, dorsolateral view of two juveniles in situ, SVL 17.8‾18.7 mm. For species from subgenus  Xenophrys, by males’ body in moderate size (SVL 31.0– 34.8 mm, n = 2),  Megophrys lushuiensis  sp. nov.differs from species with larger males:  M. aceras(55.8–62.4, n = 6),  M. ancrae(39.1–45.3, n = 8),  M. auralensis(76.7, n = 1),  M. awuh(35.7–41.1, n = 4),  M. damrei(47.7–57.1, n = 7),  M. flavipunctata( 56.9– 68.4 mm, n = 4),  M. glandulosa(76.3–81.0, n = 10),  M. himalayana(68.0–73.5, n = 6),  M. lekaguli(55.6–66.6, n = 8),  M. longipes(47, n = 1),  M. major(71.6–87.5, n = 12), M. mangshanensis(62.5, n = 1),  M. maosonensis(77, n = 1),  M. medogensis(57.2–68.7, n = 21),  M. megacephala(45.9–53.4, n = 12),  M. monticola(38.4–49.5, n = 17),  M. pachyproctus(35.3–35.7, n = 2),  M. periosa(71.3–93.8, n = 12),  M. robusta(73.5–83.1, n = 6),  M. serchhipii(36.1–46.7, n = 25),  M. shuichengensis(102.0–118.3, n = 7), and  M. takensis(47.3–53.0, n = 3). By males’ body in moderate size (SVL 31.0– 34.8 mm, n = 2),  Megophrys lushuiensis  sp. nov.differs from  M. vegrandis(vs. 27.5–30.6, n = 4 inthe latter). By vomerine teeth absent,  Megophrys lushuiensis  sp. nov.differs from  M. dzukou,  M. numhbumaeng,  M. oreocrypta,  M. oropedion,  M. pachyproctus,  M. parva,  M. zhangi, and  M. zunhebotoensis(vs. present in latter).   FIGURE 6.Photos of the tadpole CIBYN201909171 of  Megophrys lushuiensis  sp. nov.in life. A, dorsal view. B, lateral view. C, ventral view. D, dorsal view of head.   TABLE 3. Measurements (in mm) of tadpole specimens of  Megophrys lushuiensis sp. nov.    Voucher  CIBYN201909171  CIBYN201909119  CIBYN201909172  CIBYN201909175  CIBYN201909176   Gosner stage 32 36 27 27 26   BH 3.9 4.7 2.9 3.0 2.1   BL 12.1 11.3 8.4 8.8 6.8   BW 4.4 4.5 3.2 3.7 3.2   ED 1.1 1.1 0.6 0.8 0.5   ES 3.4 3.5 2.3 2.6 1.9   NE 0.8 0.8 0.6 0.6 0.6   IND 2.4 2.4 1.7 1.9 1.6   RN 2.5 2.4 1.8 2.0 1.7   SS 7.1 6.5 4.9 5.1 4.5   SU 14.4 12.4 7.7 10.3 7.5   ODW 7.6 7.7 5.7 5.6 4.1   TAL 30.7 29.8 21.6 21.3 15.7   LF 1.4 1.4 0.6 1.0 0.4   UF 1.3 1.2 0.6 0.8 0.5   TMW 2.7 2.8 0.7 1.8 1.6   TMH 3.6 3.8 2.4 2.4 1.8   TOL 42.7 41.1 30.3 29.9 23.1 For species in subgenus  Panophrys, by males’ body in moderate size (SVL 31.0– 34.8 mm, n = 2),  Megophrys lushuiensis  sp. nov.differs from species with larger males:  M. baolongensis(41.8–45.0, n = 5),  M. baolongensis(41.8–45.0, n =5),  M. binlingensis(45.1–51.0, n = 3),  M. caobangensis(34.9–38.9, n = 11),  M. caudoprocta(70.8–81.3, n = 4),  M. chishuiensis(43.4–44.1, n = 3),  M. daweimontis(34–37, n = 18),  M. hoanglienensis(43.2 ± 2.2, n = 11),  M. huangshanensis(36.0–41.6, n = 4),  M. insularis(36.8–41.2, n = 5),  M. jiangi(34.4–39.2, n = 9),  M. jingdongensis(53.0–56.5, n = 3),  M. jinggangensis(35.1–36.7, n = 2),  M. liboensis(61.6–62.9, n = 4),  M. mirabilis(55.8–61.4, n = 2),  M. obesa(35.6, n = 1),  M. omeimontis(56.0–59.5, n = 10),  M. palpebralespinosa(36–38.9, n = 2),  M. sangzhiensis(54.7, n = 1),  M. spinata(47.2–54.4, n = 18),  M. xiangnanensis(38.6–42.0, n = 11). By males larger (SVL 31.0– 34.8 mm, n = 2),  Megophrys lushuiensis  sp. nov.differs from smaller species:  M. cheni(26.2– 29.5, n = 15),  M. mufumontana(30.1–30.8, n = 2),  M. rubrimera(26.7–30.5, n = 8),  M. shimentaina(28.0–30.6, n = 12). By vomerine ridge present,  Megophrys lushuiensis  sp. nov.differs from  M. binchuanensis,  M. boettgeri,  M. kuatunensis,  M. leishanensis,  M. lishuiensis,  M. ombrophila,  M. shunghuangensis,  M. wuliangshanensis, and  M. wushanensis(vs. absent in the latter). By vomerine teeth absent,  Megophrys lushuiensis  sp. nov.differs from  M. dongguanensis,  M. fansipanensis,  M. jiulianensis,  M. nankunensis,  M. nanlingensis(vs. present in the latter). By tongue slightly notched behind,  Megophrys lushuiensis  sp. nov.differs from  M. brachykolos,  M. tuberogranulata,  M. acuta,  M. angka,  M. lini,  M. wugongensis, and  M. yangmingensis(vs. not notched behind in the latter). By toes with rudimentary webbing,  Megophrys lushuiensis  sp. nov.differs from  M. shuichengensis(vs. distinctly webbed in the latter). By tympanum about half size of eye and tibia longer than half of body length (TD/EL 0.54–0.54, TIB/ SVL 0.56–0.56, n = 2),  Megophrys lushuiensis  sp. nov.differs from  M. minor(vs. tympanum distinctly large TD/EL 0.80–0.83 and tibia relatively shorter TIB/SVL 0.46–0.48, n = 4 inthe latter).  There are twelve phylogenetically close species for the new species, of which nine species have nuclear gene data.  Megophrys lushuiensis  sp. nov.differs from  M. daweimontisby relatively smaller male body size (SVL 31.0– 34.8 mm, n = 2 vs. 34–37 mm, n = 18 inthe latter); vomerine teeth absent (vs. present), tibio-tarsal articulation reaching the middle eye when leg stretched forward (vs. reaching the tip of snout), and ventral hindlimbs semitransparent purplish with greyish white pigments (vs. reddish; Fig. 9G-L).The new species differs from  M. spinataby a horn-like tubercle on edge of upper eyelid small (vs. absent), toes with narrow fringes and rudimentary webbing (vs. fringes and webbing well developed). The new species differs from  M. sangzhiensisby eyes relatively smaller, TD/ EL 0.54, n = 2 (vs. eyes relatively larger TD/EL 0.62, n = 1), ventral hindlimbs semitransparent purplish with greyish white pigments (vs. ventral thigh and tibia orange red). The new species differs from  M. binchuanensisby dorsal skin relatively smooth (vs. smooth), tubercle on edge of upper eyelid small (vs. absent), and lateral fringes on toes narrow (vs. wide). The new species differs from  M. binlingensisby ventral hindlimbs semitransparent purplish with greyish white pigments (vs. ventral thigh and tibia dark orange red), and distinct patches scattered on middle ventral body present (vs. absent). The new species differs from  M. palpebralespinosaby dorsal skin relatively smooth (vs. rough), horn’-like tubercle on edge of upper eyelid small (vs. slightly large), vomerine teeth absent (vs. present), tongue slightly notched behind (vs. not notched), and lateral fringes on toes narrow (vs. wide). The new species differs from  M. omeimontisby vomerine ridge weak, vomerine teeth absent (vs. vomerine ridge strong, vomerine teeth present), and ventral hindlimbs semitransparent purplish with greyish white pigments (vs. ventral thigh and tibia orange red). The new species differs from  M. jingdongensisby tympanum rounded (vs. oval), vomerine teeth absent (vs. present), and toes with narrow fringes and rudimentary webbing, webbing formula: I 2–2 ¾ II 2–3¾ III 3–4¼IV 4–3V (vs. fringes and webbing well developed, webbing formula: I 1–2 II 1–3 III 2–3½IV 3½–1½ V, Fig. 9A-F). The new species differs from  M. wuliangshanensisby dorsal skin relatively smooth (vs. rough with dense granules), and lateral fringes on toes narrow (vs. absent); vomerine ridge weak (vs. absent).   TABLE 4.References for morphological characters of congeners of the genus  Megophrys.     ID  Species  Literature obtained  1   Megophrys acerasBoulenger, 1903  Bourret 1942; Munir et al.2018  2   Megophrys acutaWang, Li, and Jin, 2014  Li et al.2014  3   Megophrys ancrae Mahony, Teeling & Biju, 2013  Mahony et al.2013  4   Megophrys angka Wu, Suwannapoom, Poyarkov, Chen, Pawangkhanant, Xu, Jin, Murphy & Che, 2019  Wu et al.2019  5   Megophrys auralensis Ohler, Swan & Daltry, 2002  Ohler et al.2002  6   Megophrys awuhMahony, Kamei, Teeling & Biju, 2020 Mahony et al.2020  7   Megophrys baluensisBoulenger, 1899  Boulenger 1899 a, 1908  8   Megophrys baolongensis Ye, Fei & Xie, 2007  Ye et al.2007; Fei & Ye 2016  9   Megophrys binchuanensis Ye & Fei, 1995  Ye et al.1995; Fei & Ye 2016  10   Megophrys binlingensisJiang, Fei & Ye, 2009  Fei et al.2009  11   Megophrys boettgeriBoulenger, 1899  Boulenger 1899b; Fei et al.2009  12   Megophrys brachykolos Inger & Romer, 1961  Inger et al.1961; Fei et al.2009; Li et al.2014  13   Megophrys caobangensisNguyen, Pham, Nguyen, Luong, & Ziegler, 2020  Nguyen et al.2020  14   Megophrys carinense Boulenger, 1889  Boulenger 1908; Bourret 1942  15   Megophrys caudoprocta Shen, 1994  Shen 1994; Shen et al.2013  16   Megophrys cheni Wang & Liu, 2014  Wang et al.2014  17   Megophrys chishuiensis Xu, Li, Liu, Wei & Wang, 2020  Xu et al.2020  18   Megophrys chuannanensis Fei, Ye & Huang, 2001  Fei & Ye 2001; Fei et al.2009  19   Megophrys damreiMahony, 2011 Mahony 2011  20   Megophrys daweimontisRao & Yang, 1997  Rao & Yang 1997a; Fei & Ye 2016  21   Megophrys dongguanensisWang & Wang, 2019  Wang et al.2019a  22   Megophrys dringiInger, Stuebing & Tan, 1995  Inger et al.1995; Oberhummer et al.2014  23   Megophrys dzukouMahony, Kamei, Teeling & Biju, 2020 Mahony et al.2020  24   Megophrys edwardinae Inger, 1989  Inger et al.1989  25   Megophrys elfina Poyarkov, Duong, Orlov, Gogoleva, Vassilieva, Nguyen, Nguyen, Nguyen, Che & Mahony, 2017  Poyarkov et al.2017 ......continued on the next page   TABLE 4.(Continued)     ID  Species  Literature obtained  26   Megophrys fansipanensis Tapley, Cutajar, Mahony, Nguyen, Dau, Luong, Le, Nguyen, Nguyen, Portway, Luong & Rowley, 2018  Tapley et al.2018  27   Megophrys feae Boulenger, 1887  Boulenger 1908; Fei et al.2009  28   Megophrys feii Yang, Wang & Wang, 2018  Yang et al.2018  29   Megophrys flavipunctata Mahony, Kamei, Teeling & Biju, 2018  Mahony et al.2018  30   Megophrys gerti Ohler, 2003  Poyarkov et al.2017  31   Megophrys gigantica Liu, Hu & Yang, 1960  Liu et al.1960; Fei et al.2009  32   Megophrys glandulosa Fei, Ye & Huang, 1990  Fei et al.1990, 1992, 2009; Fei & Ye 2016  33   Megophrys hansi Ohler, 2003  Ohler 2003  34   Megophrys himalayana Mahony, Kamei, Teeling & Biju, 2018  Mahony et al.2018  35   Megophrys hoanglienensis Tapley, Cutajar, Mahony, Nguyen, Dau, Luong, Le, Nguyen, Nguyen, Portway, Luong & Rowley, 2018  Tapley et al.2018  36   Megophrys huangshanensis Fei & Ye, 2005  Fei & Ye 2005; Fei et al.2009; Fei & Ye 2016  37   Megophrys insularisWang, Liu, Lyu, Zeng & Wang, 2017  Wang et al.2017a  38   Megophrys intermedia Smith, 1921  Smith 1921  39   Megophrys jiangi Liu, Li, Wei, Xu, Li, Wang & Wu, 2020  Liu et al.2020  40   Megophrys jingdongensis Fei & Ye, 1983 Fei at al. 1983; Fei et al.2009; Fei & Ye 2016  41   Megophrys jinggangensis Wang, 2012  Wang et al.2012  42   Megophrys jiulianensis Wang, Zeng, Lyu & Wang, 2019  Wang et al.2019a  43   Megophrys kalimantanensisMunir, Hamidy, Matsui, Iskandar, Sidik& Shi- mada, 2019  Munir et al.2019  44   Megophrys kobayashii Malkmus & Matsui, 1997  Malkmus & Matsui 1997  45   Megophrys kouiMahony, Foley, Biju & Teeling, 2017 Yang 1991  46   Megophrys kuatunensis Pope, 1929  Pope 1929; Fei et al.2009; Tapley et al.2017  47   Megophrys lancip Munir, Hamidy, Farajallah & Smith, 2018  Munir et al.2018  48   Megophrys leishanensisLi, Xu, Liu, Jiang, Wei & Wang, 2019“2018”  Li et al.2018a  49   Megophrys lekaguliStuart, Chuaynkern, Chan-ard & Inger, 2006  Stuart et al.2006a  50   Megophrys liboensis Zhang, Li, Xiao, Li, Pan, Wang, Zhang & Zhou, 2017  Zhang et al.2017  51   Megophrys ligayae Taylor, 1920  Taylor 1920  52   Megophrys lini Wang & Yang, 2014  Wang et al.2014  53   Megophrys lishuiensis Wang, Liu & Jiang, 2017  Wang et al.2017b  54   Megophrys longipes Boulenger, 1886  Boulenger 1908; Bourret 1942  55   Megophrys major Boulenger, 1908  Boulenger 1908  56   Megophrys mangshanensis Fei & Ye, 1990  Fei et al.1990, 1992; Fei & Ye 2016  57   Megophrys maosonensis Bourret, 1937  Bourret, 1937; Bourret 1942  58   Megophrys medogensis Fei, Ye & Huang, 1983 Fei at al. 1983; Fei et al.2009; Fei & Ye 2016  59   Megophrys megacephala Mahony, Sengupta, Kamei & Biju, 2011  Mahony et al.2011 ......continued on the next page   TABLE 4.(Continued)     ID  Species  Literature obtained  60   Megophrys microstomaBoulenger, 1903  Fei et al.2009  61   Megophrys minor Stejneger, 1926  Stejneger 1926; Li et al.2014; Fei & Ye 2016  62   Megophrys mirabilis Lyu, Wang & Zhao, 2020  Lyu et al.2020  63   Megophrys montanaKuhl and Van Hasselt, 1822  Munir et al.2018  64   Megophrys monticolaG̹nther, 1864  Mahony et al.2018  65   Megophrys mufumontanaJ. Wang, Lyu & Y.Y. Wang, 2019  Wang et al.2019a  66   Megophrys nankiangensisLiu and Hu, 1966  Fei et al.2009  67   Megophrys nankunensisWang, Zeng&. Wang, 2019  Wang et al.2019a  68   Megophrys nanlingensisLyu, J. Wang, Liu & Y.Y. Wang, 2019  Wang et al.2019a  69   Megophrys nasuta Schlegel, 1858  Mahony et al.2018  70   Megophrys numhbumaengMahony, Kamei, Teeling & Biju, 2020 Mahony et al.2020  71   Megophrys obesa Wang, Li & Zhao, 2014  Li et al.2014  72   Megophrys ombrophila Messenger & Dahn, 2019  Messenger et al.2019  73   Megophrys omeimontis Liu, 1950  Fei et al.2009; Fei & Ye 2016  74   Megophrys oreocrypta Mahony, Kamei, Teeling & Biju, 2018  Mahony et al.2018  75   Megophrys orientalis Li, Lyu, Wang & Wang, 2020  Li et al.2020  76   Megophrys oropedion Mahony, Teeling & Biju, 2013  Mahony et al.2013  77   Megophrys pachyproctus Huang, 1981  Huang & Fei 1981; Huang et al.1998  78   Megophrys palpebralespinosa Bourret, 1937  Bourret 1942  79   Megophrys parallela Inger & Iskandar, 2005 Inger & Isk&ar 2005  80   Megophrys parva Boulenger, 1893  Boulenger 1908; Deuti et al.2017  81   Megophrys periosa Mahony, Kamei, Teeling & Biju, 2018  Mahony et al.2018  82   Megophrys popei Zhao, Yang, Chen, Chen & Wang, 2014  Zhao et al.2014  83   Megophrys platyparietusRao & Yang, 1997  Rao & Yang 1997b; Li et al., 2020  84   Megophrys robusta Boulenger, 1908  Boulenger 1908; Mahony et al.2018  85   Megophrys rubrimeraTapley, Cutajar, Mahony, Chung, Dau, Nguyen, Luong & Rowley, 2017  Tapley et al.2017  86   Megophrys sangzhiensis Jiang, Ye & Fei, 2008  Jiang et al.2008  87   Megophrys serchhipii Mathew & Sen, 2007  Mathew & Sen 2007  88   Megophrys shapingensis Liu, 1950  Liu et al.1950; Fei et al.2009  89   Megophrys shimentaina Lyu, Liu & Wang, 2020  Lyu et al.2020  90   Megophrys shuichengensis Tian & Sun, 1995  Tian & Sun 1995; Fei et al.2009  91   Megophrys shunhuangensis Wang, Deng, Liu, Wu & Liu, 2019  Wang et al.2019b  92   Megophrys spinataLiu & Hu, 1973  Fei et al.2009; Fei & Ye 2016  93   Megophrys stejnegeri Taylor, 1920  Taylor 1920  94   Megophrys synoriaStuart, Sok & Neang, 2006  Stuart et al.2006b  95   Megophrys takensisMahony, 2011 Mahony 2011 ......continued on the next page   TABLE 4.(Continued)     ID  Species  Literature obtained  96   Megophrys tuberogranulataShen, Mo & Li, 2010  Mo et al.2010  97   Megophrys vegrandis Mahony, Teeling & Biju, 2013  Mahony et al.2013  98   Megophrys wawuensisFei, Jiang & Zheng, 2001  Fei et al.2009  99   Megophrys wugongensisJ. Wang, Lyu & Y.Y. Wang, 2019  Wang et al.2019a  100   Megophrys wuliangshanensis Ye & Fei, 1995  Fei et al.2009; Fei & Ye 2016  101   Megophrys wushanensis Ye & Fei, 1995  Fei et al.2009; Fei & Ye 2016  102  Megophrys xiangnanensis Lyu, Zeng & Wang, 2020  Lyu et al.2020  103   Megophrys xianjuensis Wang, Wu, Peng, Shi, Lu & Wu, 2020  Wang et al.2020  104   Megophrys yangmingensis Lyu, Zeng & Wang, 2020  Lyu et al.2020  105   Megophrys zhangi Ye & Fei, 1992  Ye & Fei 1992; Fei et al.2009; Fei & Ye 2016  106   Megophrys zunhebotoensis Mathew & Sen, 2007  Mathew & Sen 2007 For three phylogenetically related species without nuclear gene data, the new species differs from  M. caobangensisby distinct patches scattered on middle ventral body present (vs. absent), and skins on ventral thigh around cloaca with spinous small granules (vs. absent), tympanum relatively larger (TD/ED 0.54 vs. 0.40–0.50); differs from  M. rubrimeraby vomerine teeth absent (vs. present), toes with narrow fringes (vs. wide), and finger tips rounded (vs. finger tips slightly expanded and flattened to oval pads); differs from  M. angkaby lateral fringes on toes narrow (vs. absent), and tibia relatively longer (TIB/SVL 0.56, n = 2 inthe new species vs. 0.47–0.50, n = 6 inthe latter).   Distribution and Ecology.  Megophrys lushuiensis  sp. nov.is currently known from Lushui County, Yunnan Province, China. This species is found sitting on herbs’ leaves near mountain stream under broad leaf forests at elevations 1986–2399 m( Fig. 7). Calls are rare during daytime of field work from 23 rdSeptember to 27 thSeptember 2019, when toadlets right after metamorphosis and tadpoles at Gosner stage 26 to 36 were collected.  M. glandulosawas recorded at lower elevation ( 1213 m) in Luzhang Town, Lushui County, Yunnan Province, China. Tadpoles of the new species were found almost in every small stream investigated. 3055517307 2019-09-27 WB SC Shi & WB Zhu & L Fu China Lushui County 2358 25.977066 Gutan River in Luzhang Town 1 98.79598 Nujiang Prefecture 9 360 CIBYN201909290 1 Yunnan Province holotype 3055517305 27 th September, 2019 WB SC Shi & WB Zhu & LFu Lushui County 1986 25.95118 Bashan 1 98.768906 Luzhang Town 9 360 CIBYN201909283, CIBYN201909288-289 1 paratype 3055517306 2019-09-27 WB Lushui County 2363 25.9587 Five 1 98.717995 Lushui County 9 360 CIBYN201909119 1 3055517301 2019-09-27 WB Lushui County 2363 25.9587 Five 1 98.717995 Lushui County 9 360 CIBYN201909119, CIBYN201909171-172, CIBYN201909175, CIBYN201909176 2 3055517303 2019-09-27 WB Lushui County 2363 25.9587 Five 1 98.717995 Lushui County 9 360 CIBYN201909119, CIBYN201909171-172, CIBYN201909175, CIBYN201909176 2 holotype 3055517302 2019-09-27 WB Lushui County 2363 25.9587 Five 1 98.717995 Lushui County 9 360 CIBYN201909119, CIBYN201909171-172, CIBYN201909175, CIBYN201909176 2 3055517304 2019-09-27 WB SC Shi & Zhu & L Fu Lushui County 2363 25.9587 Five 1 98.717995 Lushui County 9 360 CIBYN201909119, CIBYN201909171-172, CIBYN201909175, CIBYN201909176 2