Megophrys shunhuangensis, Wang & Deng & Liu & Wu & Liu, 2019

Wang, Lu, Deng, Xuejian, Liu, Yong, Wu, Qianqian & Liu, Zhao, 2019, A new species of the genus Megophrys (Amphibia: Anura: Megophryidae) from Hunan, China, Zootaxa 4695 (4), pp. 301-330 : 308-319

Megophrys shunhuangensis sp. nov.

Holotype: Adult male, HNNU16SH02, collected by Shengchao Shi on 28 Sep 2016, from a small stream at the edge of a mountain outside Lizhu village (500m linear distance), Shunhuang Mountain (26.441990º N, 111.013876 ºE; 882 m a.s.l.), Xinning County, Hunan Province, China.

Paratypes: Ten adults (9 males, 1 female): Two adult males, HNNU16SH08 and HNNU16SH13, collected from the same locality as the holotype by Shengchao Shi on 28 Sep 2016 ; two adult males, HNNU16SH06 and HNNU16SH07, collected by Shengchao Shi on 29 Sep 2016 in a moss-covered small stream outside Lizhu village . An adult male, HNNU18NS03, collected by Xuejian Deng on 16 Sep 2018 , from a stream beside the hillside road to Nanshan Town , Nanshan National Forest Park (26.156547ºN, 110.175120ºE; 1102 m a.s.l.), Chengbu County, Hunan Province, China; two adult males, HNNU18NS04 and HNNU18NS05, collected from the same locality as the HNNU18NS03 by Yong Liu on 18 Sep 2018 ; two adult males, HNNU18NS01 and HNNU18NS02, collected from a stream beside an abandoned canal near (1.2 km linear distance) the acquisition location of HNNU18NS03 (110.166623 ºE, 26.148937 ºN; 1233 m a.s.l.) by Qianqian Wu , Yong Liu and Lu Wang on 19 Sep 2018 . An adult female, HNNU16SH04, collected by Shengchao Shi, from the same locality as the holotype on 28 Sep 2016 ; One juvenile, HNNU16SH14, collected from the same locality as the holotype by Shengchao Shi on 28 Sep 2016 .

Diagnosis: Megophrys shunhuangensis sp. nov. is characterized by the combination of the following characters: (1) a small-sized species with 30.3–33.7mm SVL in 10 adult males and 37.6mm SVL in 1 adult females; (2) head length slightly shorter than head width (HL/HW ratio ranges from 0.81–0.89:1); (3) snout round in dorsal view, slightly projecting, sloping backward to mouth in lateral view, protruding well beyond margin of lower jaw; (4) tympanum small but distinct, rounded (TYD /ED ratio ranges from 0.48–0.67:1); (5) pupil vertical; (6) maxillary teeth present; (7) vomerine teeth absent; (8) margin of tongue smooth, oblateness, not notched behind; (9) hindlimb slender, tibio-tarsal articulation reaches forward between the nasal and tip of snout, relative finger length I <II <IV <III; (10) heels overlapping when tibia depressed against femur are held at right angles to the body axis; (11) lateral fringes absent, toes with rudimentary webbing at their bases; (12) distinct subarticular tubercles at the base of each finger; (13) a small horn-like tubercle at the middle edge of each upper eyelid; (14) dorsal skin mostly smooth with scattered granules and a few tubercles, some of which form a “two Y bottoms connected”-shaped ridge and two discontinuous dorsolateral ridges on either side of dorsal; (15) ventral surface smooth; (16) four limbs mostly smooth with longitudinal scattered granules; (17) supratympanic fold distinct and become moderately enlarged and glandular in the posterior edge; (18) dorsal skin brick red (adobe brown) with a brown triangular marking formed between the outer margin of the middle upper eyelids, a “two Y bottoms connected”-shaped light brown spots in the dorsocentral region; (19) white reticulation in the middle of lower ventral surface; (20) several dark brown crossbars on dorsal limbs, lower arm with a single dark wide stripe about half the length of it, dorsal hand has a single dark brown stripe; (21) nuptial pad present, nuptial spines tiny and densely covered in adult males, single vocal sac in males.

Holotype description (measurements in mm): HNNU16SH02, mature male, body slender and small ( SVL 33.6 mm). Head moderately large, width slightly larger than length ( HW 11.7mm, HL 10.5mm, HL/HW ratio 0.89:1); snout round in dorsal view, slightly projecting, sloping backward to mouth in lateral view, protruding well beyond margin of lower jaw; dorsal surface of snout flattened; canthus rostralis developed; loreal region vertical, a little concave; nostril positioned laterally; eye large and bright, eye diameter is 0.39 of head length and 2.14 times maximum diameter of visible portion of tympanum ( ED 4.2mm, TYD 2.0mm, HL 10.5mm); interorbital distance slightly wider than upper eyelid ( IOS 3.2mm, UEW 3.0mm); tympanum small but distinct, rounded, with upper border slightly concealed by supratympanic ridge; eye–tympanum distance shorter than tympanum diameter ( TYE 1.7mm, TYE / TYD ratio 0.86:1); pupil vertically rhomboid; ovoid choanal slit at base of maxillary shelves; vomerine ridges and vomerine teeth absent; maxillary teeth present; margin of tongue smooth, oblateness, not notched behind, a discontinuous linear line on the posterior edge of the tongue.

Forelimbs slender, measuring 46% of SVL, hand length slightly shorter than forearm length ( LAHL 15.4mm, FAL 8.3mm, HAL 8.2mm); fingers without lateral fringes; interdigital webbing absent; subarticular tubercle at base of each finger; metacarpal tubercle two, inner one elliptical and moderate, outer one smaller and flat; finger tips rounded with translucent bulge below them, slightly expanded relative to digit widths, terminal grooves absent. Hindlimbs slender, measuring 50% of SVL, thighs slightly shorter than tibia, and slightly longer than feet ( TL 17.0mm, THL 15.2mm, FOL 14.1mm); tibio-tarsal articulation reaches forward between the nasal and tip of snout; heels overlapping when tibia depressed against femur are held at right angles to the body axis; toes long, without lateral fringes, relative toe lengths I <II <V < III <IV; toe tips rounded with translucent dilated below them, basically equal width to toe, terminal grooves absent; hands without webbing, toes with rudimentary webbing at their bases relative finger length I <II <IV < III; inner metatarsal tubercle small and elliptical, outer metatarsal tubercles absent; tarsal fold absent.

Skin of dorsal surfaces of body, limbs, and dorsal and lateral surfaces of head primarily smooth with scattered granules and a few tubercles; several tubercles on the upper eyelid, presence of a horn-like tubercle in the middle edge of the eyelid; tympanum smooth, slightly concave, with borders slightly raised; canthus ridges developed; supratympanic fold distinct, narrow anteriorly gradually expanding beyond posterior edge of tympanum to become moderately enlarged and glandular, curving from posterior edge of orbits, along the upper part of the tympanum, curving down abruptly at posterior tympanum border to scapula; dorsolateral ridges absent; weakly developed tubercles form a “two Y bottoms connected”-shaped ridge and two short discontinuous dorsolateral ridges on either side of the “two Y bottoms connected”-shaped ridges; several large tubercles scattered on flanks and lateral parts of the body; small scattered granules on dorsal surface of limbs, 3–4 stripes on the back of the thighs and shanks, each stripe has a short row of smaller tubercles. Ventral surfaces of head, body and limbs smooth; pair of small, white and weakly raised pectoral glands on chest; femoral glands large, flat, almost on posterior surface of middle thighs; ventral side covered with white small granules; ventral thighs and upper arm mostly smooth, with several small white granules below them ( Fig. 4 ).

Holotype measurements (in mm): SVL 33.6, HL 10.5, HW 11.7, SL 4.38, INS 3.61, IOS 3.21, UEW 3.01, ED 4.18, TYD 1.95, TYE 1.67, SN 2.16, LAHL 15.36, FAL 8.32, HAL 8.23, LAD 3.07, HLL 52.62, THL 15.21, TL 16.99, TFL 22.47, FL 14.1, TW 4.25.

Coloration of holotype in life: Dorsal and lateral surfaces of head and body primarily brick red with lighter brown tones; lighter brown stripe on the dorsal snout; a brown with light-edged triangular marking is formed between the middle of the outer margin of the upper eyelids; front of snout taupe brown with closely orange red granules; narrow circle of olive spots around the eyes; dark brown alternate with light brown blotch extends from the posterior border of eye to the posterior tympanum; lower half of supratympanic ridges dark brown, upper half brick red, between them with a long white stripe from the posterior edge of orbits, along the upper part of the tympanum, curving down abruptly at posterior tympanum border to scapula; dark brown rectangle blotch distinct between the lower orbital border and the upper lip; dark brown stripe on upper lip extends from nostril to rear of upper jaw, dark brown stripes on both sides of the lower lip margin; a lighter brown “two Y bottoms connected”-shaped marking on anterior dorsum, lighter brown, discontinuous stripe present on each side of dorsolateral; dorsal surfaces of limbs primarily brick red with lighter brown blotches; forearm has a single dark brown wide stripe, about half the length of them; dorsal hand has a single dark brown stripe; dorsal surfaces of hindlimbs with distinct brown spindle transverse blotches; dorsal surface of outer three fingers with dark brown blotches, dorsal toes distribute light brown blotches. A distinct dividing line is formed between the ventral and the dorsal include limbs by a white stripe; ventral surface brown, white reticulation blotches in the middle of down ventral, dark brown on both sides of the abdomen; throat primarily lighter brown, possessing a single dark brown with pale yellow narrow edge wide streak almost throughout throat the down the midline; chest and anterior abdomen with dark orange blotches on otherwise brown background; ventral surfaces of limbs primarily orange red scattered with white spots; subarticular tubercles, inner and outer metacarpal tubercles and inner metatarsal tubercle are all fluorescent red; ventral surfaces of tarsi and feet brown, hands lighter brown; pectoral and femoral glands milky white ( Fig. 4 ).

Coloration of holotype in preservation: Preservation caused increased contrast in specimen compared to life. Majority of dorsal and lateral surfaces of the head, body, limbs lighter brown; the triangular marking between the upper eyelids become dark brown and the light-edged milky white; closely orange red granules front of snout become white; front of snout and from posterior orbital border to the posterior tympanum become dark brown; the lower margin of the temporal fold is dark brown and the upper margin is light brown with a white stripe in the middle; large spots on both sides of the upper and lower lip and between the lower orbital border and the upper lip are also dark brown; “two Y bottoms connected”-shaped marking on anterior dorsum and discontinuous dorsolateral stripe become brown; the single wide stripe in the forearm and the stripe in the dorsal hand also dark brown; the spindle transverse blotches on dorsal surfaces of hindlimbs also brown; dorsal surface of outer three fingers with dark brown blotches, dorsal toes distribute brown blotches. Throat region, chest and anterior part of abdomen primarily lighter brown; milky white reticulation blotches in the middle of down ventral; dark brown spots on both sides of the abdomen with distinct demarcation from the dorsal; area surrounding vent with dark brown blotches; pectoral and femoral glands milky white; ventral surfaces of limbs with lighter brown comminution mottling; ventral surfaces of feet brown, hands almost milky white with lighter brown blotches; tips of fingers and toes, subarticular tubercles, inner and outer metacarpal tubercles and inner metatarsal tubercle are all milky white ( Fig. 5 ).

Variation: See Table 2 for measurements and body proportions within the type series consisting of nine adult males, one adult female, and a juvenile. The paratypes resemble the holotype for most morphological characters with some exceptions: some individuals dorsal skin adobe brown in life; the whole body has distinct red tubercles in life (HNNU16SH04, HNNU16SH07, HNNU16SH13, HNNU16SH14), tubercles of HNNU16SH13 are especially distinct in the inguinal and also especially obvious in the head of HNNU16SH14, the only juvenile; tubercles on the dorsal of HNNU16SH04, HNNU18NS01-03 became particularly marked in preservation; two V-shaped ridge on anterior dorsum in HNNU18NS01; the dorsal of HNNU16SH08 has two V-shaped marking; most specimens had distinct “two Y bottoms connected”-shaped marking on anterior dorsum; HNNU16SH06, HNNU16SH07 and HNNU18NS02 has a single oval streak almost in the midline of dorsal snout; the three sides of the triangle between the upper eyelids are inward concave in HNNU16SH04 and HNNU16SH13; two dark brown rectangle blotch distinct between the lower orbital border and the upper lip in HNNU16SH04; supratympanic ridges conceal up to ~20% of upper tympanum in HNNU16SH13, HNNU18NS04; dark brown streak in the throat almost invisible in HNNU16SH06, HNNU18NS04 and HNNU18NS05, not obvious in HNNU16SH07, HNNU16SH13 and HN- NU18NS02; discontinuous dark brown spots on both sides of the abdomen in HNNU16SH04; pectoral glands on some individuals marginally down of chest.

Secondary sexual characteristics: Females are slightly larger than males in body size. Males: nuptial pads on Finger I present and weakly raised, covered with black micro-asperities, water drop-shaped, covering most of dorsal surface of Finger I; nuptial pad on Finger II, strip shape, positioned on base of digit on inner dorsal side, from the base of proximal phalange to the base of fingertip, almost 2/3 the length of the Finger II; nuptial spines tiny and densely covered; single vocal sac present, moderately large internal vocal slits present on floor of mouth near rear of mandible on each side; forearms and tibias shorter and thicker than females (males LAD/FAL ratio 0.33–0.40:1, TW/TL ratio 0.23–0.25:1; females LAD/FAL ratio 0.30, TW/TL ratio 0.20:1). Female: nuptial pads, vocal sac, vocal slits, all absent.

Etymology: This specific epithet shunhuangensis is a Latinized toponymic adjective that refers to the type locality: Shunhuang Mountains, Xinning County, Hunan Province, China. On the other hand, it is to honor Shun, a Chinese ancient Saint Emperor.

Suggested vernacular name: Shunhuang horned frog (English) and 舜皇角蟾 (Chinese).

Distribution: Currently M. shunhuangensis sp. nov. is known from the vicinity of Lizhu village (within 500m of the town) and Nanshan National Forest Park, Hunan Province, China.

Ecology: This species inhabit subtropical mountain forest at elevation between 882 to 1233 m a.s.l., in southern Hunan and northern Guangxi province. We observed 12 individuals in total (10 male, 1 female, 1 juvenile), all through active search methods. No eggs were observed despite search efforts. All specimens were collected on September. Provenance of finds respective ranged between 882m and 1102–1233 m. Seven males from Shunhuang Mountain were found calling on or under dense bush leaves in a small mountain stream at temperature 16.4°C, one male was found calling on banana ( Musa . sp) leaves about 1 meter above ground ( Fig. 6 ). Nanshan specimens were found on foggy nights between 0 and 2 am, and the temperature was about 6–10°C, five individuals were found in small clearances washed by mountain streams, surrounded by subtropical evergreen broadleaved forests, among them, two specimens were found on a leaf by the stream ( Fig. 7 ).

Morphological comparisons. Megophrys shunhuangensis sp. nov. can be distinguished from all other congeners found in Megophrys . Here we compare the undescribed species with all known 85 congeners known as of April 2019. Including 80 kinds from Amphibiaweb and M. elfina Poyarkov et al. 2017 , M. lancip Munir et al. 2018 , M. lishuiensis ( Wang et al. 2017) , M. maosonensis Bourret 1937 , M. leishanensis . Comparative data and the diagnosis of subgenus from the literature: Rao & Yang 1997; Li & Wang 2008; Chen et al. 2017; Mahony et al. 2017; Fei et al. 2009; Messenger and Dahn 2019; Li et al. 2018; Mahony et al. 2018; Yang et al. 2018; Li et al. 2014; Wang et al. 2017; Tapley et al. 2017; Poryakov et al. 2017 and AmphibiaWeb 2019. All specimens examined for morphometric analysis are listed in Table 3 .

Megophrys shunhuangensis sp. nov. can be distinguished from the species in the subgenus Atympanophrys : M. gigantica Liu et al. 1960 , M. nankiangensis Liu et al. 1966 , M. shapingensis Liu 1950 and M. wawuensis Fei et al. 2001 by the presence of a tympanum and a horn-like tubercle in the eyelid. In Atympanophrys the tympanum is absent or indistinct and palpebral projections absent.

Megophrys shunhuangensis sp. nov. can be distinguished from the species in the subgenus Brachytarsophrys : M. carinense ( Boulenger 1889) , M. chuannanensis ( Fei and Ye 2001) , M. feae Boulenger 1887 , M. intermedia Smith 1921 and M. popei by the heels overlapping and smaller adult male size, SVL 30.3–33.6mm (versus SVL> 79.1 mm). In Brachytarsophrys , when hindlimbs are flexed, heels are not overlapping.

Megophrys shunhuangensis sp. nov. can be distinguished from the species in the subgenus Ophryophryne : M. gerti ( Ohler 2003) , M. hansi ( Ohler 2003) , M. koui Kou 1985 , M. microstoma ( Boulenger 1903) and M. synoria ( Stuart et al. 2006) by the presence of a small and indistinct horn-like tubercle at middle the edge of the upper eyelid (versus with a relatively prominent and elongated tubercle). Differs from M. elfina by the smooth with small tubercles dorsal skin (versus dorsal skin shagreened). Differs from M. synoria by having a smaller adult male size, SVL 30.3–33.6 mm (versus SVL 38.2–53.7 mm) and further from all species in the subgenus Ophryophryne due to the presence of maxillary teeth (versus maxillary teeth absent).

Megophrys shunhuangensis sp. nov. can be distinguished from the species in the subgenus Megophrys : M. lancip , M. montana ( Kuhl and Van Hasselt 1822) and M. parallela ( Inger and Iskandar 2005) by the distribution and tubercles on the outer edge of the upper eyelids short, short tubercle on the tip of snout absent. Subgenus Megophrys is endemic to Indonesia with long horn-like tubercles on the outer edge of the upper eyelids and a short tubercle on the tip of snout.

Megophrys shunhuangensis sp. nov. differs from 18 species: M. aceras Bourret 1903 , M. auralensis Ohler et al. 2002 , M. damrei Mahony 2011 , M. dringi Inger et al. 1995 , M. flavipunctata Mahony et al. 2018 , M. glandulosa Fei et al. 1990 , M. himalayana Mahony et al. 2018 , M. lekaguli Stuart et al. 2006 , M. longipes Boulenger 1886 , M. major Boulenger 1908 , M. mangshanensis , M. maosonensis , M. medogensis Fei et al. 1983 , M. megacephala Mahony et al. 2011 , M. oreocrypta Mahony et al. 2018 , M. periosa Mahony et al. 2018 , M. robusta Boulenger 1908 and M. takensis Mahony et al. 2011 by the smaller adult male size, SVL 30.3-33.6mm (versus SVL> 45.9 mm). Differs from 9 species: M. ancrae Mahony et al. 2013 , M. monticola (Günther 1864) , M. oropedion Mahony et al. 2013 , M. pachyproctus Huang and Fei 1981 , M. serchhipii ( Mathew and Sen 2007) , M. zhangi Ye and Fei 1992 and M. zunhebotoensis ( Mathew and Sen 2007) by the absence of vomerine teeth (versus vomerine teeth present). Differs from M. feii Yang et al. 2018 and M. vegrandis Mahony et al. 2013 by the nuptial pad present, nuptial spines tiny and densely covered in adult males (versus nuptial pad absent). Differs from M. cf. parva by the distinct subarticular tubercles (versus subarticular tubercles indistinct or absent). Differs from M. huangshanensis by a slender hindlimb, tibio-tarsal articulation reaching forward between the nasal and tip of snout, no notched tongue and heels overlapping (versus shorter hindlimb, tibio-tarsal articulation reaching forward between the posterior border of eye and tympanum; tongue feebly notched; heels not meeting).

Megophrys shunhuangensis sp. nov. is placed by molecular evidence ( Fig. 2 ) into the subgenus Panophrys . Within this subgenus it differs from 10 species: M. baolongensis Ye et al. 2007 , M. binlingensis , M. caudoprocta , M. hoanglienensis Tapley et al. 2018 , M. jingdongensis Fei et al. 1983 , M. liboensis , M. omeimontis Liu 1950 , M. sangzhiensis , M. shuichengensis Tian and Sun 1995 and M. spinata Liu and Hu 1973 by smaller adult male size, SVL 30.3–33.6mm (versus SVL> 45.9 mm). It dffers from M. acuta , M. cheni , M. jinggangensis and M. ombrophila by nuptial pad present (versus nuptial pad absent). Differs from five species: M. daweimontis Rao and Yang 1997 , M. fansipanensis Tapley et al. 2018 , M. insularis , M. latidactyla and M. rubrimera by the absence of vomerine teeth (versus vomerine teeth present). Differs from M. palpebralespinosa , M. tuberogranulata and M. wuliangshanensis Fei et al. 1995 by the distinct subarticular tubercles (versus subarticular tubercles absent). Differs from M. binchuanensis Ye et al. 1995 , M. boettgeri , M. lini and M. wushanensis by the absence of lateral fringes ( M. binchuanensis , M. boettgeri and M. lini with widely lateral fringes; M. wushanensis with widely lateral fringes in male, without in female). Differs from M. kuatunensis Pope 1929 and M. minor Stejneger 1926 by the not notched tongue (versus tongue feebly notched). Differs from M. brachykolos , M. lishuiensis and M. obesa by heels overlapping when thighs are positioned at right angles to the body ( M. brachykolos and M. obesa , versus heels not meeting; in M. lishuiensis , heels not meeting or just meeting).

Two members of the subgenus Panophrys , Megophrys leishanensis and M. acuta , in the molecular phylogeny are very close to the new species. The new species differs from M. leishanensis by the smooth with small tubercles dorsal skin, tibio-tarsal articulation reaches between eye and snout, relative finger length I <II <IV <III and with a single dark brown wide stripe about half the length of the forearm (versus dorsal skin rough; tibio-tarsal articulation reaches between tympanum and eye; relative finger lengths II <I <IV <III; without wide stripe about half the length of the forearm or just narrow stripe). Differs from M. acuta by the smooth with small tubercles dorsal skin, overlapping heels, tibio-tarsal articulation reaches between eye and snout, lacking lateral fringes on toes and nuptial pad present, nuptial spines tiny and densely covered in adult males (versus dorsal skin smooth; heels not meeting; tibio-tarsal articulation reaches in ocular region; lateral fringes narrow; nuptial pad weak, no nuptial spines in adult males); for the SVL of M. shunhuangensis sp. nov. and M. acuta , M. acuta is slightly smaller.

Other species that in the molecular phylogeny are placed close to the new species, or have low genetic divergences to it, are Megophrys baolongensis , M. boettgeri , M. caudoprocta , M. cheni , M. huangshanensis , M. jinggangensis , M. kuatunensis , M. liboensis , M. minor , M. tuberogranulata , M. wushanensis . In morphological comparisons, the new species differs from M. baolongensis by the smaller adult male size (SVL 30.3–33.6mm), not notched tongue and tibio-tarsal articulation reaches between eye and snout (versus SVL 41.8–45.0mm; tongue feebly notched; tibio-tarsal articulation reaches in ocular region). Differs from M. boettgeri by the not notched tongue, tibio-tarsal articulation reaches between eye and snout and lacking lateral fringes on toes (versus tongue feebly notched; tibio-tarsal articulation reaches in ocular region; lateral fringes on toes wide). Differs from M. caudoprocta by the absent vomerine teeth, tibio-tarsal articulation reaching between eye and snout, overlapping heels and lacking lateral fringes on toes (versus vomerine teeth present; tibio-tarsal articulation reaching in ocular region; heels not meeting or just meeting; lateral fringes on toes narrow or lacking). Differs from M. cheni by the not notched tongue, lacking lateral fringes on toes, distinct subarticular tubercles and nuptial pad present (versus tongue notched; lateral fringes on toes wide; subarticular tubercles indistinct or absent; nuptial pad absent). Differs from M. huangshanensis by the not notched tongue, tibio-tarsal articulation reaches between eye and snout and overlapping heels (versus tongue feebly notched; tibio-tarsal articulation reaches between eye and tympanum; heels not meeting or just meeting). Differs from M. jinggangensis by the smooth with small tubercles dorsal skin, absent vomerine teeth, lacking lateral fringes on toes and nuptial pad present (versus dorsal skin rough; vomerine teeth present; lateral fringes on toes narrow; nuptial pad absent). Differs from M. kuatunensis by the smooth with small tubercles dorsal skin, not notched tongue, tibio-tarsal articulation reaching between eye and snout and overlapping heels (versus dorsal skin smooth; tongue feebly notched; tibio-tarsal articulation reaching between eye and tympanum; heels not meeting or just meeting). Differs from M. liboensis by the smaller adult male size (SVL 30.3–33.6mm), smooth with small tubercles dorsal skin, absent vomerine teeth, not notched tongue, tibio-tarsal articulation reaching between eye and snout, lacking lateral fringes on toes and distinct subarticular tubercles (versus SVL 34.7–67.7mm; dorsal skin smooth; vomerine teeth present; tongue feebly notched; tibio-tarsal articulation reaching in ocular region; lateral fringes on toes narrow; subarticular tubercles indistinct or absent). Differs from M. minor by the not notched tongue and tibio-tarsal articulation reaching between eye and snout (versus tongue feebly notched; tibio-tarsal articulation reaching in ocular region). Differs from M. tuberogranulata by the smooth with small tubercles dorsal skin, overlapping heels and distinct subarticular tubercles (versus dorsal skin rough; heels not meeting or just meeting; subarticular tubercles indistinct or absent). Differs from M. wushanensis by the smooth with small tubercles dorsal skin, tibio-tarsal articulation reaches between eye and snout and lacking lateral fringes on toes (versus dorsal skin smooth; tibio-tarsal articulation reaches in ocular region; widely lateral fringes in male, without in female).

For five members of the subgenus Panophrys , no molecular data are available: Megophrys daweimontis , M. fansipanensis , M. hoanglienensis , M. latidactyla and M. shuichengensis . In morphological comparisons, the new species differs from M. daweimontis by the smooth with small tubercles dorsal skin, absent vomerine teeth, not notched tongue and distinct subarticular tubercles (versus dorsal skin rough; vomerine teeth present; tongue feebly notched; subarticular tubercles absent). Differs from M. fansipanensis and M. hoanglienensis by the absent vomerine teeth, not notched tongue and distinct subarticular tubercles (versus vomerine teeth present; tongue feebly notched and subarticular tubercles absent). Differs from M. latidactyla by the smooth with small tubercles dorsal skin, absent vomerine teeth and absent lateral fringes (versus dorsal skin rough; vomerine teeth present; lateral fringes widely). Differs from M. shuichengensis by the smaller adult male size (SVL 30.3–33.6mm), not notched tongue, tibio-tarsal articulation reaches between eye and snout, absent lateral fringes, distinct subarticular tubercles and present nuptial pad (versus SVL 102.0– 118.3mm; tongue feebly notched; tibio-tarsal articulation reaches in ocular region; lateral fringes widely; subarticular tubercles absent; nuptial pad absent).

Figures

FIGURE 2. Bayesian inference tree derived from partial DNA sequences of the mitochondrial 12S rRNA + 16S rRNA gene. Numbers after slashes are bootstrap support for maximum likelihood (1000 replicates) analyses (>50 retained) and numbers before slash indicate Bayesian posterior probabilities (>0.50 retained). The symbol “-” represents bootstrap value below 50.

FIGURE 4. Megophrys shunhuangensis sp. nov. holotype: adult male (HNNU16SH02: SVL 33.6 mm) in life: A. dorsal view; B. ventral view; E. lateral view showing dark wide stripe in the forearm; F. ventral view of hand; G. ventral view of foot. Paratype: adult female (HNNU16SH04: SVL 37.6 mm) in life C. dorsal view; D. ventral view.

FIGURE 5. Megophrys shunhuangensis sp. nov. holotype: adult male (HNNU16SH02: SVL 33.6 mm) in preservation: A. dorsal view; B. ventral view; E. ventral view of hand; F. ventral view of foot; G. nuptial pads on the first and second fingers; H. profile view of head; I. discontinuous linear line on the posterior edge of the tongue. Paratype: adult female (HNNU16SH04: SVL 37.6 mm) in preservation C. dorsal view; D. ventral view.

FIGURE 6. Habitats of M. shunhuangensis sp. nov. in the type locality, a small stream at the edge of Shunhuang Mountain.

FIGURE 7. Habitats of M. shunhuangensis sp. nov. in the type locality, a small-clearance washed by mountain streams.

Tables

TABLE 3. Diagnostic characters separating M. shunhuangensis sp. nov. from all known 63 recognized species of subgenus Xenophrys and subgenus Panophrys. Comparative data were obtained from the literature (Rao & Yang, 1997; Li & Wang, 2008; Fei et al., 2009; Li et al., 2014; Chen et al., 2016; Wang et al., 2017 a; Tapley et al., 2017; Mahony et al., 2017; Mahony et al., 2018; Yang et al., 2018; Messenger et al., 2019; Li et al., 2019).

	SVL males	females	Dorsal skin texture: rough (++), smooth with small tuber- cles (+), smooth (-)	Vomerine teeth: present (+), or absent (-)	Tongue: notched (++), feebly notched (+), or not notched (-)	Tibio-tarsal articula- tion reaches forward: be- tween eye and snout(+++), in ocular region(++), be- tween eye and tympanum(+), after(-)	Heels: overlap- ping (+), not meeting or just meeting (-)	Lateral fringes on toes: wide (++), narrow (+), lacking (-)	Subarticular tubercles: distinct (+), indistinct or absent (-)	Nuptial pad: present (+), ab- sent(-)
M. shunhuangensis sp. nov.	30.34–33.65	37.6	+	-	-	+++	+	-	+	+
M. aceras	55.8–62.4	67–86	+	+	/	++	-	+	/	/
M. acuta	27.1–33.0	28.1–33.6	-	-	-	++	-	+	+	-
M. ancrae	39.1–45.3	48.9	+	+	+	/	/	-	-	+
M. auralensis	76.7	/	+	-	-	/	/	+	-	+
M. baolongensis	41.8–45.0	/	+	-	+	++	+	-	+	+
M. binchuanensis	32.0–36.0	40.2–42.5	-	-	+ or -	++	-	++	+	+
M. binlingensis	45.1–51.0	/	+	-	+	++	+	+	+	+
M. boettgeri	34.5–37.8	39.7–46.8	+	-	+	++	+	++	+	+
M. brachykolos	33.7–39.3	33.9–45.9	+	-	-	-	-	-	+	+
M. caudoprocta	70.8–81.3	77.8	+	+	-	++	-	+ or -	+	+
M. cheni	26.2–29.5	31.8–34.1	+	-	++	+++	+	++	-	-
M. damrei	57.1	69.1	-	+	++	/	/	-	-	+
M. daweimontis	34–37	40–46	-	+	+	+++	/	-	-	/
M. dringi	42.8–46.7	55	+	-	/	/	/	-	-	-
M. fansipanensis	30.9–44.3	41.7–42.5	+	+	+	/	/	-	-	+
M. feii	24.5–25.1	28.2–28.9	++	-	+	/	+	++	-	-
M. flavipunctata	56.9–68.4	68.0–74.6	-	+	++	/	/	+	-	+
M. glandulosa	76.3–81.0	76.5–99.5	-	+	+	+++	+	++	-	+
M. himalayana	68.0–73.5	83.9	+	+	/	/	/	-	-	+

TABLE 3. Diagnostic characters separating M. shunhuangensis sp. nov. from all known 63 recognized species of subgenus Xenophrys and subgenus Panophrys . Comparative data were obtained from the literature ( Rao & Yang, 1997; Li & Wang, 2008; Fei et al., 2009; Li et al., 2014; Chen et al., 2016; Wang et al., 2017 a; Tapley et al., 2017; Mahony et al., 2017; Mahony et al., 2018; Yang et al., 2018; Messenger et al., 2019; Li et al., 2019).

......continued on the next page