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Megophrys (Xenophrys) awuh sp. nov. Holotype. Adult male (BNHS 6070 [field tag SDBDU 2007.163] [Figure 5]) from Star Lake (25.464444, 94.703889, 2220 m asl.), Reguri village, Meluri circle, Phek district, Nagaland state, Northeast India, collected by RGK and VK on 5 July 2007. Paratypes. Adult female (BNHS 6069 [field tag SDBDU 2007.161]) and adult male (BNHS 6071 [field tag SDBDU 2007.162]) collected along with holotype. Referred specimens. Adult male (SDBDU 2007.111 [Figure 6 (a)]) from about 1 km from Dzükou tourist entry office (25.571667, 94.120278, ca. 1650 m asl.), off highway NH 39, Jakhama village, Kohima district, Nagaland state, NEI, collected by RGK and VK on 24 June 2007; adult male (SDBDU 2007.079 [Figure 6 (e – f)]) and adult female (SDBDU 2007.080 [Figure 6 (b – d)]) from the river Dzütsürü (25.613611, 93.969444, 2010 m asl.), Dzüleke village, Sechu-Zubza circle, Kohima district, Nagaland state, NEI, collected by RGK and VK on 18 June 2007; tissue sample without associated specimen (SDBDU 2007.192) from Aradurah Forest, above New Ministers ' Hill (25.641389, 94.094167, 1980 m asl.), Kohima town, Kohima Sadar circle, Kohima district, Nagaland state, NEI, collected by RGK and SDB on 10 July 2007. Holotype description (measurements in mm). Mature male (SVL 37.2) (Figure 5). Head moderately large, as long as wide (HW 13.0, HL 13.0, IFE 6.3, IBE 10.3); snout bluntly pointed in dorsal view, obtusely protruding in lateral view, rostral appendage absent (Figure 5 (c)); loreal region acute, concave; canthus rostralis angular; dorsal region of snout slightly concave; eye larger than maximum tympanum diameter, and equal to snout length (EL 4.7, TYD 2.8, SL 4.7); eye – tympanum distance (TYE 2.2) shorter than maximum tympanum diameter; tympanum distinctly oval shaped, upper border not concealed by supratympanic ridge; pupil oval in preservation, orientated vertically; nostril oval, orientated laterally, closer to eye than to snout (EN 2.2, NS 2.6); internarial distance greater than eyelid width, and equal to narrowest point between upper eyelids (IN 3.9, UEW 3.7, IUE 3.9); pineal ocellus not visible externally; vomerine ridges weak, forming long narrow ridges without swollen posterior end, orientated acutely, extending from near anterior choanae border, terminating posterior to choanae, posterior ends equidistant from each other and choanae; vomerine teeth absent; maxillary teeth present; tongue large, covers almost entire floor of mouth, cordiform (with weak notch posteriorly), no medial lingual process. Forelimbs moderately long, thin (Figure 5 (a – b)); forearms slightly enlarged relative to upper forelimbs, forearm shorter than hand length (FAL 9.1, HAL 10.1); fingers moderately long, thin, lateral dermal fringes absent (Figure 5 (d)), finger length formula I = II – b)); thigh shorter than shank, and foot (TL 16.7, SHL 18.3, FOL 17.1); toes long, rounded, distinct lateral dermal fringes absent (Figure 5 (e)), relative toe lengths I Skin of dorsal surfaces of body, limbs, and dorsal and lateral surfaces of head smooth (Figure 5 (a)); tympanum smooth with borders weakly raised; ridge or tubercle on outer edge of upper eyelids absent; supratympanic ridge narrow anteriorly, expanding posterior to tympanum to become moderately large and glandular, extending from orbit in a straight line to upper tympanum border, broadly curving obliquely downward along posterior border of tympanum, terminating above forearm insertion on each side; flanks with numerous small to large pustular tubercles (Figure 5 (a)); dorsolateral ridges thick, well defined, extending from behind supratympanic ridge to above groin on each side; weak, ' X ' -shaped ridge present on dorsum, its two sides extending posteriorly from above anterior border of tympanum and terminating beyond level of axilla on parietoscapular region, and posterior ends join to dorsolateral ridges on mid dorsum (Figure 5 (a)); weak tubercles and short, obliquely transverse ridges present on dorsal surfaces of thighs, shanks and forearms. Gular region, chest, abdomen and ventral surfaces of limbs all smooth; pectoral glands medium-sized, flat, positioned on level of axilla (Figure 5 (b)); femoral glands moderately large, slightly raised, positioned equidistant from knee and cloaca on posterior surface of thighs; black dermal asperities restricted to tympanic regions, posterior upper eyelids and posterior-lateral portion of head near supratympanic ridges, absent from all remaining surfaces. Colour: In preservation (Figure 5): Dorsal and lateral surfaces of head and body primarily light brownish-grey; solid dark brown triangular marking on dorsal surface of head between eyes; longitudinal brown stripe on mid-dorsal snout, extending from medial anterior border of triangular marking to tip of snout; dorsal ridges and flank tubercles light brownish-cream, some with small bordering dark brown spots and speckling; front of snout and lateral canthus rostralis dark brown; wide vertical dark brown bar below eyes; dark brown blotch extends from posterior canthus, through tympanum to posterior supratympanic ridge on each side; supratympanic ridges light yellowish-cream; dorsal surfaces of forelimbs and hindlimbs primarily light yellowish-grey; two dark brown blotches on anterior lateral surface of forearms; dorsal surface of outer three fingers with dark brown blotches; lateral surfaces of thighs and shanks with dark brown spots and blotches. Throat, chest and abdomen primarily pale grey, lightening to creamishyellow on lateral and posterior surfaces of abdomen and ventral surfaces of forelimbs, thighs and shanks; throat with faint dark brown longitudinal blotches, those most defined extending from rear axis of mandibles to base of forelimbs on each side, and third on midgular region extending onto anterior chest; dark brown blotches form broken stripe laterally along abdomen on both sides; area surrounding vent and posterior surfaces of thighs dark brown; ventral surfaces of tarsus and feet dark brown with contrasting light grey callous ridges on toes; light grey oval patch on region where inner metatarsal tubercle would be on other species; forelimbs and hands ventrally mottled creamishyellow and light greyish-brown, ventral surfaces of digits light grey; pectoral and femoral glands creamish-white. In life: No images in life taken for this specimen. Variation. See Table 2 for morphometric variation within the specimen series of four adult males and two adult females. Paratypes and the referred specimen resemble the holotype for most morphological characters with some exceptions: finger length formula typically I = II ' V ' -shaped marking on anterior dorsum surrounding parietoscapular ridges; SDBDU 2007.111 possesses small tubercle on outer upper eyelid, SDBDU 2007.079 and SDBDU 2007.080 have short ridge on upper eyelid (Figure 6 (a)); parietoscapular and dorsal ridge configuration differs from holotype as follows: SDBDU 2007.111 has faintly raised, longitudinal ' >-< ' configuration (Figure 6 (a)), SDBDU 2007.079 and SDBDU 2007.080 possess only ' V ' -shaped parietoscapular ridges (Figure 6 (c,e)); BNHS 6069, BNHS 6071 and SDBDU 2007.080 have no observed dermal asperities, SDBDU 2007.111 has white dermal asperities found only on middorsum, increasing in density posteriorly to above vent, on SDBDU 2007.079, asperities only found on posterior surface of upper eyelids; SDBDU 2007.111, SDBDU 2007.079 and SDBDU 2007.080 are mid to light brown dorsally; on BNHS 6069 and BNHS 6071, triangular brown marking on dorsal surface of the head is faint, ventrolateral dark brown blotches are smaller and less dense, and do not form broken stripe laterally along abdomen on both sides, and ventral shanks and thighs with dark brown, dense speckling; BNHS 6069 has small dark brown spots spread over entire abdomen. Secondary sexual characters. Males: weakly raised nuptial pads covered with brown or black micro-asperities, covering most of dorsal surface of base of Finger I, narrowing distally, extending onto base of distal phalange on inner dorsal side; on Finger II, nuptial pad small, oval shaped, widest proximally, extending just onto basal phalange on inner dorsal side; external vocal sac indistinct; internal vocal slits present near rear of lower mandible; forearms slightly enlarged relative to upper forelimbs; fleshy projection above cloaca absent. Females: nuptial pads, vocal slits, cloacal projection all absent; forearms not enlarged relative to upper forelimbs; both females contained moderately large, pigmentless ova. Morphological comparisons. Megophrys awuh sp. nov. (adult males, N = 4; adult females, N = 2) differs from the following species that have not yet been assigned to a subgenus or species group through molecular analyses: from M. damrei and M. shuichengensis by smaller adult size, male SVL 35.7 – 41.1 mm, female SVL 43.7 – 48.0 mm (vs. M. damrei: adult male SVL 47.7 – 57.1 mm, N = 7, adult female SVL 69.1 mm, N = 1 [Mahony 2011; Neang et al. 2013]; M. shuichengensis: adult male SVL 102.0 – 118.3 mm, N = 8, adult female SVL 99.8 – 115.6 mm, N = 7 [Tian et al. 2000]); from M. feii by larger adult size, male SVL 35.7 – 41.1 mm, female SVL 43.7 – 48.0 mm (vs. adult male SVL 24.3 – 25.1 mm, N = 4, adult female SVL 28.2 – 28.9 mm, N = 2), lateral dermal fringes on toes absent (vs. present), and nuptial pads on fingers of males present (vs. absent) (Yang et al. 2018). Megophrys awuh sp. nov. differs from M. parva s.s. by larger tympanum relative to eye length, TYD/ED 55.8 – 72.7%, N = 6 (vs. TYD/ED 40.0 – 55.0%, N = 4), longer relative forearm length, FAL/SVL 24.5 – 26.3%, N = 6 (vs. FAL/SVL 20.9 – 23.9%, N = 4), longer relative shank length, SHL/SVL 49.2 – 54.9%, N = 6 (vs. SHL/SVL 42.8 – 48.2%, N = 4), fingertips not expanded relative to finger width (vs. slightly expanded relative to finger width). Megophrys awuh sp. nov. differs from species molecularly assigned to the M. (X.) megacephala SG based on the following combinations of characters: differs from M. ancrae by finger and toe tips not expanded relative to digit width (vs. digit tips distinctly expanded terminally), small raised bump or short ridge on outer upper eyelids sometimes present (vs. short, sharp horn-like tubercle on outer upper eyelids present), circummarginal patch of demal asperities ventrally on mandible of males absent (vs. present on seven of eight males examined), and typically narrower snout, IFE/ HW 45.3 – 49.6 %, N = 6 (vs. 49.6 – 57.1%, N = 10); differs from M. megacephala by smaller adult male size, SVL 35.7 – 41.1 mm (vs. SVL 45.9 – 53.4 mm, N = 7), and considerably narrower head, HW/ SVL 31.5 – 36.7 %, N = 6 (vs. HW/ SVL 40.2 – 45.1 %, N = 12); differs from M. oropedion by slightly longer relative shank length, SHL/ SVL 49.2 – 54.9 %, N = 6 (vs. SHL/ SVL 41.1 – 48.4 %, N = 12), vomerine teeth usually absent, five of six specimens (vs. usually present, nine of 11 specimens examined), dermal asperities, if present, sparsely distributed, may be found on tympanic region, mid to posterior dorsum, and posterior upper eyelids (vs. extensively distributed on most dorsal, and some ventral surfaces of head, body and limbs); differs from M. serchhipii by FIIL≤FIVL (vs. FIVL– 54.9%, N = 6 (vs. FOL/ SVL 44.3 ± 2.2%, range 39.5 – 48.7%, N = 32); differs from M. zunhebotoensis by larger adult male size, SVL 35.7 – 41.1 mm (vs. 28.4 – 33.9 mm, N = 22), typically larger tympanum diameter relative to eye diameter ratio, TYD/ED (mean±SD) 60.1 ± 2.8%, range 55.7 – 63.6%, N = 6 (vs. TYD/ ED 50.5 ± 3.9%, range 42.9 – 58.5%, N = 26), tympanum oval (vs. circular). For a comparison with the two new species described below, see the comparison sections for those species. Etymology. The specific epithet 'awuh' is treated as a noun in apposition taken from the Pochury language, meaning ' frog '. The Pochury tribe are the native inhabitants of Meluri circle in Nagaland state, within which the type locality lies. Suggested common name. We suggest ' Naga Hills Horned Frog ' for this species based on its currently known distribution. Distribution. Known only from the vicinity of Kohima and Star Lake in southern and eastern Nagaland state, respectively, at elevations between 1650 and 2220 m asl. (Figure 2). Considering the wide elevational range inhabited, the geographic distribution of this species is expected to be more extensive than currently reported. The proximity of the collection localities relative to political borders indicates that Megophrys awuh sp. nov. will most likely be found in neighbouring Manipur state (NEI) and Sagaing Region (western Myanmar) at similar elevations. Habitat and natural history. All males were calling when collected, and both females contained mature ova indicating that the breeding season includes at least the mid monsoon months of June and July. The type specimens were collected from a primary forest with lush undergrowth, patches of closed canopy, hill streams, and where there were not any significant signs of human activities except perhaps occasional and regulated hunting by the local Pochurys. All specimens were collected during daylight hours (12:00 – 16:00 h). The holotype was found calling from within grasses on the edge of a spring about a metre deep and three metres diameter; the male paratype was found calling beneath grasses half submerged in a puddle in the forest. The female paratype was collected from the edge of the same pond as the holotype. The referred specimen SDBDU 2007.111 was collected from ground level in secondary forest adjacent to highway NH 39 on a night with light rain, about 10 m away from a seasonal stream. SDBDU 2007.079 and SDBDU 2007.080 were collected at ~20:00 h from vegetation within 10 m of the river Dzütsürü in a mature secondary forest. The Aradurah Forest tissue sample was collected from a juvenile (SVL ~ 12 mm) caught on a rainy night at ~19:30 h on the ground under a bush. The bush was situated about four metres away from a seasonal hill stream. A few males were heard calling but possibly due to the moderately heavy rain the calls were not prolonged enough for the individuals to be located. Remarks. Molecular phylogenetic analyses consistently resolved Megophrys awuh sp. nov. (adult male SVL 35.7 – 41.1 mm) as the sister taxon of the smaller M. zunhebotoensis (male SVL 28.4 – 33.9 mm) in all analyses that included mtDNA (Figures 1 & S1; Mahony et al. 2017, figs. 1 & 3 [as 'M. sp. [2 & 3] ']; Mahony et al. 2018, fig. 4 [as 'M. sp. 5 ']). Molecular dating analyses suggested that these sister taxa diverged from their most recent common ancestor at ca. 3.8 MY (Mahony et al. 2017, fig. 1).

Mahony, Stephen, Kamei, Rachunliu G., Teeling, Emma C., Biju, S. D. (2020): Taxonomic review of the Asian Horned Frogs (Amphibia: Megophrys Kuhl & Van Hasselt) of Northeast India and Bangladesh previously misidentified as M. parva (Boulenger), with descriptions of three new species. Journal of Natural History 54 (1-4): 119-194, DOI: 10.1080/00222933.2020.1736679, URL: http://dx.doi.org/10.1080/00222933.2020.1736679

Megophrys (Xenophrys) himalayana sp. nov. (Figures 20 & 21; Table 1) Holotype. Adult male (BNHS 6050 [field no. SDBDU 2009.1227]: Figures 20, & 21A, F & H), from Elephant village (27°4'56.52"N, 92°34'50.22"E, 370 m asl.), West Kameng district, Arunachal Pradesh state, Northeast India, collected by members of the Systematics Lab, University of Delhi on either 0 4 or 0 5 August 2009. Paratypes. Four adult males (BNHS 6051–6054 [field nos. SDBDU 2009.1206 – 1209]: Figure 21C, D, F & H), from Elephant village collected along with the holotype by members of the Systematics Lab, University of Delhi between 0 4 and 0 5 August 2009. Referred specimens. Adult male (SDBDU 2009.787: Figure 21E & G), and adult female (SDBDU 2009.750: Figure 21B & G), from Rengging village (28°8'32.52"N, 95°16'18.72"E, 410 m asl.), East Siang district, Arunachal Pradesh state, Northeast India, collected by Systematics Lab members between 24 and 25 July 2009. Holotype description (measurements in mm). Mature male (SVL 72.1) (Figures 20, & 21A, F & H). Head moderately large, as wide as long (HW 27.0, HL 26.9, IFE 13.3, IBE 21.4); snout bluntly pointed in dorsal view, obtusely protruding beyond mandible in lateral view, without rostral appendage (Figure 20C); loreal region acute, concave; canthus rostralis angular; dorsal surface of snout very slightly concave; eye diameter more than twice maximum diameter of visible portion of tympanum, slightly shorter than snout length (EL 9.0, TYD 4.0, SL 9.6); eye–tympanum distance (TYE 5.8) approximately two times diameter of visible portion of tympanum; tympanum oval-shaped, oblique, with upper border concealed by supratympanic ridge (Figure 20C); pupil vertically elliptical; nostril positioned laterally, closer to eye than to snout tip (EN 4.0, NS 5.6); internarial distance greater than eyelid width and narrowest point between upper eyelids (IN 8.7, UEW 7.8, IUE 7.8); pineal ocellus not visible externally; vomerine ridge present, medium sized, ovoid, moderately raised, orientated acutely, positioned between to slightly posterior to choanae, equidistant from choanae and each other; vomerine teeth short; maxillary teeth present; tongue not observed due to fixation of jaw and in interest of preventing potential damage to jaws by forcing open mouth wide enough for examination. Forelimbs long, thin (Figure 20A & B), forearm moderately enlarged relative to upper forelimb, and shorter than hand length (FAL 16.7, HAL 19.0); fingers long, narrow, without lateral fringes (Figure 20D), finger length formula IV Skin of dorsal and lateral surfaces of head, body, and limbs primarily smooth, sparsely covered with small, weakly defined granules; tympanum with borders raised relative to surrounding region; outer edge of upper eyelids with a broad distinctly pointed bump; supratympanic ridges narrow anteriorly, gradually expanding beyond posterior edge of tympanum to become moderately enlarged and glandular, extending from orbits, curving downward abruptly at posterior upper border of tympanum, terminating above forelimb insertions; flanks sparsely covered with small to medium sized unevenly scattered pustular tubercles; dorsolateral ridges thin, weakly developed, extending from behind supratympanic ridges to ~95% trunk length; V-shaped parietoscapular ridge weakly developed, composed primarily of single rows of closely spaced asperities; dorsal surfaces of forearms, thighs and shanks with short transverse ridges; posterior thighs with small scattered pustular tubercles; gular region, chest, abdomen and ventral surfaces of limbs smooth; pectoral glands small, weakly raised, on chest level with axilla; femoral glands moderately large, flat, on posterior surface of thighs, slightly closer to knees than to cloaca; small white dermal asperities form broad dense band circummarginally on gular region, moderately dense on upper lips, tympanic region (except tympanum), supratympanic ridge and posterior dorsal surface of upper eyelids, sparse on posterior dorsal surface of head and anteriormost dorsum of body increasing in density posteriorly, present along all dorsal ridges, absent from all remaining surfaces. Colouration: In preservative (Figure 20): Dorsal and lateral surfaces of body, and dorsal surface of head primarily mid brown; light-edged, darker brown incomplete triangular marking between eyes; dorsum with distinct darker brown X-shaped marking on centre and dorsolateral longitudinal stripe on each side; tubercles on flanks primarily dark brown, many with small lighter tips; large dark brown blotch covers tympanic region; dark brown bar extends vertically from lower orbital border; lateral surfaces of canthus rostralis and front of snout dark brown; upper lip without contrasting light longitudinal stripe; dorsal and lateral surfaces of forelimbs and hindlimbs primarily mid brown; dark brown blotches on dorsolateral surface of forearms; dorsal surface of fingers with dark brown blotches; dorsal surfaces of thighs and shanks with distinct dark brown transverse crossbars; throat, chest and anterior abdomen primarily dark brown with darker brown blotches and mottling, throat with creamish-white spots and blotches along edge of mandible; posterior half of abdomen primarily creamish-white; ventral surfaces of forelimbs, thighs and shanks mid brown with lighter yellowish-brown mottling, ventral tarsi dark brown; area surrounding vent and posterior surfaces of thighs dark brown; ventral surfaces of feet and hands greyish-brown; pectoral and femoral glands creamish-white. In life (Figure 21A & F): Dorsal surfaces of head, body and hindlimbs orangish-brown, distinctly paler on flanks and forelimbs; ventral surfaces generally lighter, with brown blotches distinct, with distinct wide brown longitudinal striped on gular region; iris dark reddish-orange. Variation. Refer to Table 1 for morphometric variation within the type series and referred specimens, consisting of six adult males and a female. Paratypes and referred specimens generally resemble the holotype for most morphological characters with the following exceptions: Head width/head length ratio varies considerably between individuals (HW/HL 94.3–106.3%); dorsolateral ridges typically weak to moderately well developed, always extending>65% of trunk length; posterior edge of tongue weakly bifurcate on three examined specimens (SDBDU 2009.750, BNHS 6051, BNHS 6053); typically only upper 10–25% of tympanum concealed by supratympanic ridge; parietoscapular-sacral ridges vary considerably in configuration, i.e., “>–– – Secondary sexual characters. Males: nuptial pads present, weakly raised, covered with black microasperities, covering most of dorsal surface of Finger I; nuptial pad on Finger II medium sized, oval, positioned on base of digit on inner dorsal side, extending onto base of proximal phalange; external vocal sac indistinct; moderately large internal vocal slits present on floor of mouth near rear of mandible on each side; forearms enlarged relative to upper forelimbs. Female: ova not pigmented; nuptial pads, vocal sac, vocal slits, enlarged forearms, all absent. Morphological comparison. Megophrys himalayana sp. nov. (adult males, N =7, adult female, N =1) differs from M. monticola and M. zhangi by its larger adult body size, male SVL 68.0– 73.5 mm, female SVL 83.9 mm (vs. male SVL 38.2–49.5 mm, N =17, female SVL 40.5–56.1 mm, N =6; male SVL 32.5–37.2 mm, N =3, respectively); differs from M. robusta by its smaller adult male size, SVL 68.0– 73.5 mm (vs. male SVL 73.5–83.1 mm, N =6), black dermal asperities on ventral surface of thighs and posterior abdomen of males absent (vs. of six adult males examined, asperities present on ventral surface of thighs of all, and on abdomen of five); differs from M. medogensis by its slightly larger adult male body size, SVL 68.0– 73.5 mm (vs. male SVL 57.2–68.0 mm, N =16), iris colour in life maroon to red (vs. light golden [Zhao et al. 2005; Li et al. 2010; Fei et al. 2010, 2012]). From MMC members, Megophrys himalayana sp. nov. differs from Megophrys flavipunctata sp. nov. by toes rounded, not flattened, lateral fringes absent (vs. toes dorsoventrally flattened, with or without narrow lateral dermal fringes), forearms of adult males moderately enlarged relative to upper arms (vs. not enlarged relative to upper arms); differs from Megophrys oreocrypta sp. nov. and M. major s.s. by dark spots associated with flank tubercles typically present (vs. absent), further from Megophrys oreocrypta sp. nov. by light portion of upper lip not extending beyond nostril (vs. light upper lip stripe extends anteriorly beyond nostril), further from M. major s.s. by webbing on Toe IV of males not extending beyond basal articulation, i.e., 4IV 4 (vs. webbing more extensive on males, i.e., 3.2IV3.2 to 3.6IV3.6), toe tips not expanded relative adjacent toe width (vs. expanded); from M. mangshanensis by larger adult body size, male SVL 68.0– 73.5 mm, female SVL 83.9 mm (vs. male SVL 62.5 mm, N =1, female SVL 73.0 mm, N =1). For comparisons with additional species covered in this study, refer to relevant morphological comparison sections. Systematic position. This taxon represents M. cf. major 5 (OTU 13) in the molecular analyses, and “ M. cf. major [4]” in Mahony et al. (2017). The systematic position of Megophrys himalayana sp. nov. within the MMC is currently unclear. Concatenated gene trees of mt+nuDNA or mtDNA only (Figures 2 & 4; Appendix I, Table 3; Appendix II, Figures 1, 2 & 5), and gene coalescence analysis of the unphased nuDNA-only dataset (Figure 5A; Appendix I, Table 3) placed this species as sister taxon to M. cf. major 4 (described as a new species, below), whereas, gene coalescence analysis of the phased nuDNA-only dataset (Figure 5B) placed this species as sister taxon to a clade comprising Megophrys flavipunctata sp. nov. and M. cf. major 4 (described as a new species, below). Refer to Appendix I, Table 6 for uncorrected p -distances for the 16S rRNA gene between Megophrys himalayana sp. nov. and other MMSG species. Etymology. The species epithet “ himalayana ” is a toponym in reference to the known distribution range of this species that is restricted to the southern Himalayas. Suggested common name: Himalayan Horned Frog. Distribution. The type series and referred specimens were collected from two widely separated localities in the state of Arunachal Pradesh, indicating an Indian distribution that ranges at least from East Siang district, west to West Kameng district, at low elevation (375–410 m asl.) (Figure 8A). A more extensive sampling is necessary to define the east-west limits of this species’ geographic range. Habitat and natural history. All specimens were collected after dusk. Animals were found typically perched on the rocky banks of small (~ 1–3 m wide), presumably temporary/seasonal, fast flowing mountain streams bordered by dense mature secondary/primary forest. This species probably favours smaller streams– –at the type locality specimens were collected from one of the small tributaries of the large Sessa stream (Figure 9E), but were not found on the banks of the Sessa stream itself. The two smallest male specimens, BNHS 6052 with no internal vocal slits and BNHS 6054 with very small internal vocal slits, had enlarged testes, well developed nuptial pads and enlarged forearms, indicating that males of this species begin reaching sexual maturity at around SVL 68.0– 68.5 mm. Although all males appeared to be in breeding condition, no vocalisations were heard at the collection localities. Furthermore, the female contained relatively under-developed ova in its ovaries indicative that the breeding season might begin sometime in late July or early August, towards the end of the monsoon season.

Megophrys (Xenophrys) monticola (Günther, 1864) (Figures 6 & 7; Table 1) Xenophrys monticola [partim] Günther 1864:415 + pl. xxvi, fig. H, In: The Reptiles of British India. The Ray Society, London: 452 pp. + xxvii + pl. 1–26. [?] Xenophrys katabhako Deuti et al. 2017:23, 27–35, In: Nomenclatural puzzle in early Xenophrys nomina (Anura, Megophryidae) solved with description of two new species from India (Darjeeling hills and Sikkim). Alytes, 34: 20–48. [?] Xenophrys sanu Deuti et al. 2017:23, 27–35, In: Nomenclatural puzzle in early Xenophrys nomina (Anura, Megophryidae) solved with description of two new species from India (Darjeeling hills and Sikkim). Alytes, 34: 20–48. Lectotype of Xenophrys monticola. Adult female (BMNH 1947.2.25.13 [R.R. [18]60.3.19.1336]: Figure 6), from “ Sikkim ” state, Northeast India, collected by the Schlagintweit brothers (presumably Hermann), some time between 14 April and 15 August 1855 (Schlagintweit et al. 1861). Lectotype designation by Mahony et al. (2017). Paralectotype of Xenophrys monticola (non M. monticola s.s.). Adult male (BMNH [18]53.8.12.52), from “Khasya” [Khasi Hills], Meghalaya state, Northeast India, presented by Sir William Jackson Hooker (BMNH Specimen Catalogue), likely collected by his son Sir Joseph Dalton Hooker between June and September 1850 (Hooker 1854). Paralectotype designation by Mahony et al. (2017). Examined referred specimens. “High-elevation” mitochondrial haplotype (2180–2220 m asl.): three adult males (SDBDU 2011.1046: Figure 7H; SDBDU 2011.1047: Figure 7C, E & H; SDBDU 2011.1048: Figure 7H) and one adult female (SDBDU 2011.1049: Figure 7F & H), from Senchal Wildlife Sanctuary (26°59'38"N, 88°18'0"E, 2220 m asl.), Rambi (East Range), Darjeeling Sadar sub-division, Darjeeling district, West Bengal state, Northeast India, collected by Systematics Lab members on 0 4 June 2011; three adult males (SDBDU 2011.429 – 431) from Senchal Wildlife Sanctuary (same locality details as above), collected by SDB and RGK on 20 May 2011; one adult male (SDBDU 2011.1029) and one adult female (SDBDU 2011.1030: Figure 7A & G), from Sukhiapokhri-Manebhanjan road (26°59'50.04"N, 88°9'58.56"E, 2180 m asl.), Darjeeling Sadar sub-division, Darjeeling district, West Bengal state, Northeast India, collected by Systematics Lab members on 0 3 June 2011; one unsexed juvenile (SDBDU 2011.1418), from Bagora Range 8th mile road marker (26°56'44"N, 88°18'00"E, 2180 m asl.), Kurseong sub-division, Darjeeling district, West Bengal state, Northeast India, collected by SDB and RGK on 20 May 2011. “Mid-elevation” mitochondrial haplotype (880–1135 m asl.): two adult males (SDBDU 2011.419 & SDBDU 2011.420), from Ghoramara area (26°53'19"N, 88°23'53"E, 1110 m asl.), Mahananda Wildlife Sanctuary, Latpanchar, Kurseong sub-division, Darjeeling district, West Bengal state, Northeast India, collected by SDB and RGK on 19 May 2011; one adult male (SDBDU 2011.418) from Stream 1 (26°54'34"N, 88°23'52"E, 1030 m asl.), Mahananda Wildlife Sanctuary, Latpanchar, Kurseong sub-division, Darjeeling district, West Bengal state, Northeast India, collected by SDB and RGK on 19 May 2011; three adult males (SDBDU 2011.1070 – 1072) from Mahananda Wildlife Sanctuary (26°53'4.5"N, 88°23'2.46"E, 1080 m asl.), Latpanchar, Kurseong sub-division, Darjeeling district, West Bengal state, Northeast India, collected by Systematics Lab members on 0 7 June 2011; two adult males (SDBDU 2011.1056; SDBDU 2011.1066: Figure 7D) and one adult female (SDBDU 2011.1065: Figure 7D), from Latpanchar town (26°54'34.2"N, 88°23'54.54"E, 1135 m asl.), Kurseong sub-division, Darjeeling district, West Bengal state, Northeast India, collected by Systematics Lab members on 0 5 and 0 6 June 2011; one adult male (SDBDU 2011.1423: Figure 7B), from Makaibari Tea Estate (26°56'44"N, 88°18'0"E, 880 m asl.), Kurseong sub-division, Darjeeling district, West Bengal state, Northeast India, collected by SDB and RGK on 21 May 2011. Not assigned to molecular haplotype: adult female lectotype (BMNH 1947.2.25.13); two adult females (BMNH [18]72.4.17C; BMNH [18]72.4.17E), from “ Sikkim ” state, Northeast India, collected by T.C. Jerdon, before 1870; one adult male (BMNH [18]72.4.17G), from “Darjeeling” district, West Bengal state, Northeast India, collected by T.C. Jerdon, before 1870; one adult female (ZSIC 9650), from “Darjeeling” district, West Bengal state, Northeast India, collected by J. Gammie. Provisionally referred specimen. One adult male (BMNH [18]72.4.17I), from “Darjeeling” district, West Bengal state, Northeast India, collected by T.C. Jerdon, before 1870. Lectotype description (measurements in mm). Mature female, SVL 40.5 (Figure 6). General preservation condition poor, dehydrated. Head large, as wide as long (HW 16.2, HL 16.2, IFE 7.2, IBE 11.8); snout rounded in dorsal view, obtuse in lateral view, protruding beyond mandible (Figure 6C); rostral appendage absent; loreal region acute and concave with well developed canthus rostralis; dorsal surface of snout concave; eye twice as large as tympanum (EL 5.6, TYD 2.8), slightly longer than snout (SL 5.4); eye–tympanum distance (TYE 2.6) slightly less than tympanum diameter; tympanum oval, obliquely orientated with upper ~20% concealed by supratympanic ridge (Figure 6C); pupil indistinct; nostril positioned laterally with raised posterior rim, equidistant from snout tip and anterior edge of eye (EN 2.7, SN 2.8) [presumed artefact of dehydration]; inter-narial distance slightly wider than space between upper eyelids and upper eyelid width (IN 4.9, IUE 4.7, UEW 4.0); pineal ocellus not visible externally; vomerine ridges small, ovoid, acutely angled, positioned equidistant from choanae and each other, situated between to slightly posterior to choanae; vomerine teeth absent; maxillary teeth present; tongue weakly notched posteriorly, medial lingual process absent. Forelimbs moderately long, thin (Figure 6A & B), forearms not enlarged relative to upper forelimbs, slightly shorter than hand length (FAL 9.5, HAL 9.7); fingers long, narrow (Figure 6D), finger length formula IColouration: In preservative (Figure 6A & B): Dorsal surface of head, body and limbs plain brown; faint darker brown triangular marking between eyes; supratympanic ridges bicoloured, upper half white, lower half dark brown; wide slightly oblique dark brown bar on upper lip below orbits (Figure 6C); gular region and chest dark brown, with dark and light blotches along edges of gular region; dark brown of chest fades posteriorly on abdomen, with some small dark brown blotches on ventrolateral surfaces of flanks; ventral thighs and shanks primarily light with darker blotches laterally; outer tarsi with continuous dark brown blotch from feet to base of shanks (Figure 6E); area surrounding cloaca dark brown; forelimbs brown above with two dark brown transverse stripes on forearms, ventrally light brown; ventral surface of hands and feet faded brown (Figure 6D & E); pectoral and femoral glands lighter than surrounding surfaces. In life: Colouration not originally documented. Description of referred specimen SDBDU 2011.1047 (measurements in mm). Mature male (SVL 40.7) (Figure 7C, E & H). Head moderately large, wider than long (HW 15.5, HL 14.9, IFE 7.7, IBE 12.1); snout bluntly pointed in dorsal view, obtusely protruding in lateral view, without rostral appendage (Figure 7C); loreal region acute, concave; canthus rostralis angular; dorsal surface of snout concave; eyes more than twice as long as maximum diameter of naked portion of tympanum, and shorter than snout length (EL 5.2, TYD 2.4, SL 5.7); eyetympanum distance (TYE 2.7) slightly longer than maximum diameter of visible portion of tympanum; tympanum oval-shaped and slightly oblique, with upper border concealed by supratympanic ridge (Figure 7C); pupils in life oval, horizontally orientated when dilated; nostrils positioned laterally, positioned closer to eyes than to snout tip (EN 2.0, NS 3.3); internarial distance equal to narrowest point between upper eyelids, and greater than eyelid width (IN 4.9, IUE 4.9, UEW 4.3); pineal ocellus not visible externally; vomerine ridges well developed, narrow, weakly raised, orientated acutely, positioned posterior to choanae, closer to choanae than to each other; vomerine teeth small; maxillary teeth present; tongue large, appears rounded posteriorly without notch, medial lingual process absent. Forelimbs moderately long, thin (Figure 7C & E), forearms moderately enlarged relative to upper forelimbs, shorter than hand length (FAL 8.4, HAL 11.4); fingers moderately long, narrow, without lateral fringes, finger length formula I=II Colouration: In preservative (Figure 7E & H): Dorsal and lateral surfaces of head, body and limbs primarily dark greyish-brown; light-edged, solid dark brown triangular marking between eyes; distinct dark brown X-shaped marking on dorsum; flank tubercles dark brown with white tips; front of snout and lateral canthus rostralis dark brown; wide vertical dark brown bar below eyes; dark brown blotch extends from posterior canthus through tympanum to posterior supratympanic ridge; two dark brown blotches on anterior lateral surface of forearms; dorsal surface of Finger III with dark brown blotches; lateral surfaces of thighs and shanks with dark brown spots and blotches; throat, chest, anterior abdomen and ventral surfaces of forelimbs, thighs and shanks primarily light brown, suffused with yellow on lateral and posterior abdomen and limbs; some small dark brown spots and blotches on abdomen; area surrounding cloaca and posterior surfaces of thighs dark brown; ventral surfaces of tarsi and feet dark brown with contrasting light grey on toes; hands ventrally mottled yellowish-white and light brown, ventral surfaces of digits light grey; femoral glands creamish-white. In life (Figure 7C): Markings as described in preservative, but general dorsal colouration light olive green with dorsal ridges, flank tubercles and larger granules suffused with orange; iris colour golden-brown; throat and chest dark brown fading to white posteriorly on abdomen; groin and inner thighs deep reddish-orange. Variation. See Table 1 for morphometric variation between the lectotype and referred specimens consisting of 17 adult males, seven adult females, and one juvenile. The remaining referred specimens resemble the lectotype and referred specimen (described above) for most morphological characters, with some exceptions: relative finger length formula varies considerably between I (”, “>––|”, “>- Secondary sexual characters. Males: weak to moderately raised nuptial pads present, covered with black micro-asperities, covering most of dorsal surface of Finger I, narrowing distally, extending to base of distal phalange on inner dorsal side; nuptial pad on Finger II small to medium sized, widest proximally, usually extending to mid-proximal phalange on inner side; large subgular external vocal sac distinct as loose skin on some specimens; internal vocal slits present near rear of mandible; forearms slightly to moderately enlarged relative to upper forelimbs. Females: mature ova without pigmented poles (diameter Morphological comparison. Megophrys monticola (adult males N =17 [excluding the provisionally referred specimen], adult females N =7) differs from M. mangshanensis by absence of distinct white upper lip stripe (vs. present). For comparisons with subsequent species covered in this paper, refer to relevant morphological comparison sections for those species. Systematic position. Megophrys monticola was found to be the sister taxon to all remaining species in the MMSG (Mahony et al. 2017; Figures 2, 3 & 4). In Darjeeling, this species has two comparatively deeply divergent mitochondrial haplotypes (Appendix I, Table 6; Appendix II, Figures 1, 5, & 7). However, nuclear diversity observed between these populations consisted of pSNPs only (Figures 3 & 5; Appendix II, Figures 3, 4, & 6) indicating recent introgression between mid- (880–1135 m asl.) and high-elevation (2180–2220 m asl.) populations. The mid-elevation populations share an identical mitochondrial haplotype with a specimen referred to the species M. zhangi by Chen et al. (2017), collected from the vicinity of its type locality (Appendix II, Figure 5). However, the RAG1 sequence of this specimen differs significantly from the Darjeeling populations (figure not provided). Further molecular sampling of intervening populations is necessary to identify the biological processes involved, and the taxonomic significance of the shared mitochondrial DNA between these species. Megophrys monticola (as redefined above) differs from the type specimens of M. zhangi by adult male size (see Morphological comparison section of M. zhangi), so our curr, Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on pages 16-30, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/2610202, {"references":["Gunther, A. (1864) The Reptiles of British India. The Ray Society, London, 452 + XXVII pp., 26 pls.","Deuti, K., Grosjean, S., Nicolas, V., Vasudevan, K. & Ohler, A. 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Neotype for Xenophrys gigas Jerdon, 1870 and Megalophrys major Boulenger, 1908 (by present designation). Adult male (BMNH 1947.2.24.93 [rr. BMNH [18]72.4.17.399]: Figure 14), from Khasi Hills, Meghalaya state [previously “Darjeeling”––see Remarks section], Northeast India, collected by T.C. Jerdon, collection date ~1870. Examined specimens. Adult male (BMNH 1947.2.24.93), neotype; four adult males (BMNH 1947.2.24.94 [rr. BMNH [18]72.4.17.400]; BMNH 1947.2.24.95 [rr. BMNH [18]72.4.17.401]; BMNH 1947.2.24.96 [rr. BMNH [18]72.4.17.402]; BMNH 1947.2.24.98 [rr. BMNH [18]72.4.17.404]), two adult females (BMNH 1947.2.24.92 [rr. BMNH [18]72.4.17.398]; BMNH 1947.2.24.97 [rr. BMNH [18]72.4.17.403]), one unsexed subadult (BMNH 1947.2.24.99 [rr. BMNH [18]72.4.17.405]), and one juvenile (BMNH 1947.2.25.1 [rr. BMNH [18]72.4.17.406]), details as per the neotype; four adult males (ZSIC 9744; ZSIC 9756; BMNH 1908.4.8.4 [ex. ZSIC 9743]; BMNH 1908.4.8.5 [ex. ZSIC 9736]), one adult female (ZSIC 9746), and one unsexed juvenile (BMNH 1908.4.8.6 [ex. ZSIC 9760]) from “Cherrapunji [=Sohra]”, East Khasi Hills district, Meghalaya state, Northeast India, collected by J.H. Bourne, collection date unknown; two adult males (SDBDU 2007.229; SDBDU 2007.230), from the Terunhü River, Kenrunhü (25°55'4"N, 94°5'24"E, 510 m asl.), New Sendenyu Village, Tseminyu sub-division, Kohima district, Nagaland, Northeast India, collected by RGK on 0 3 October 2007; one adult female (CES 18901 [field number RGK 0073]: Figure 15A & B), from Raenghkucpaeng duithuak (duithuak = stream, in local Rongmei [Nruangmei] language), near St. Joseph School (24°51'40"N, 93°38'33"E, 470 m asl.), Khumhzi II village, Luangmai (Noney) district (erstwhile [before 2016] in Tamenglong district), Manipur state, Northeast India, collected by RGK on 23 May 2013; adult male (CES 18902 [field number RGK 0088]: Figure 15C & D), from Mpeih baek duithuak (24°51'0"N, 93°37'23"E, 450 m asl.), Khumhzi village, Luangmai district, Manipur state, Northeast India, collected by RGK on 26 May 2013; adult male (CES 18903 [field number RGK 0089]: Figure 15E & F), from near Khangchiuluanh (Khongjaron) Church (24°58'1"N, 93°29'53"E, 1255 m asl.), Khangchiuluanh village, Tamenglong district, Manipur state, Northeast India, collected by RGK on 27 May 2013. Neotype description (measurements in mm). Mature male (SVL 75.0) (Figure 14). Head moderately large, longer than wide (HW 29.0, HL 29.7, IFE 12.6, IBE 20.9); snout bluntly pointed in dorsal view, obtusely protruding in lateral view, without rostral appendage (Figure 14C); loreal region acute, concave; canthus rostralis angular; dorsal surface of snout very slightly concave; eye length ~2.5 times as long as maximum diameter of visible portion of tympanum, and shorter than snout length (EL 9.2, TYD 3.7, SL 10.6); eye–tympanum distance (TYE 6.3) longer than diameter of visible portion of tympanum; tympanum oval-shaped, obliquely orientated with upper ~20% concealed by supratympanic ridge (Figure 14C); pupil vertically elliptical; nostril positioned laterally, closer to eye than to snout tip (EN 4.2, NS 6.5); internarial distance greater than upper eyelid width, and equal to narrowest point between upper eyelids (IN 9.4, UEW 6.0, IUE 9.4); pineal ocellus not visible externally; vomerine ridges present, medium sized, ovoid, moderately raised, orientated acutely, positioned between to slightly posterior to choanae, slightly closer to choanae than to each other; vomerine teeth short; maxillary teeth present; tongue moderately large, with small notch posteriorly, medial lingual process absent. Forelimbs long, thin (Figure 14A & B), forearms moderately enlarged relative to upper forelimbs, and shorter than hand length (FAL 18.4, HAL 18.7); fingers long, narrow, without lateral fringes (Figure 14D), finger length formula IV–Colouration: In preservative (Figure 14): Dorsal and lateral surfaces of body, and dorsal and lateral surfaces of head primarily brown; light-edged, faintly darker brown triangular marking between eyes; no X-, Y-, or V-shaped markings on dorsum; tubercles primarily white on posterior flanks only; lateral surfaces of head below supratympanic ridges and canthus rostralis dark brown with cream-white stripe on upper lip, extending from nostril to rear of jaw, bordered below by thin dark brown stripe; outer edges of eyelids dark brown; dorsal and lateral surfaces of forelimbs and hindlimbs primarily mid brown; three dark brown blotches on dorsolateral surface of forearms; dorsal surface of outer three fingers with faint brown blotches; dorsal surfaces of hindlimbs without distinct transverse crossbars; lateral surfaces of thighs and shanks with dark brown spots and blotches; throat and chest primarily plain mid brown with few cream-white spots along edge of gular region, light-edged wide dark brown stripe extending from posterior edge of mandible onto base of forearms; abdomen, ventral surfaces of forelimbs, thighs, shanks, and dorsal surfaces of tarsi and feet immaculate pale brown; some small dark brown spots and blotches laterally along lower flanks on both sides bordering lower-most tubercles; area surrounding vent and posterior surfaces of thighs primarily mid to dark brown with small cream-white tubercles; ventral surfaces of tarsi and feet pale greyish-brown; hands ventrally pale greyish-brown; pectoral and femoral glands creamish-white. In life: Not documented for neotype. Refer to Figure 15 for examples of colour in life of referred specimens. Variation. Refer to Table 1 for morphometric variation within the referred specimen series, consisting of 12 adult males, three adult females, two subadults, and a juvenile. Referred specimens generally resemble neotype for most morphological characters with the following exceptions: Webbing between digits varies slightly between individuals and sexes, i.e., webbing formula on Toe IV: males from 3.2IV3.2 to 3.6IV3.6, females 4IV 4.3; lateral fringes on toes vary from moderately weak from bases to tips on all toes, to almost completely absent on some individuals; relative finger lengths seem to vary between populations, IVSecondary sexual characters. Males: nuptial pads present, weakly raised, covered with brown/black microasperities on freshly collected specimens (appear as microgranules on old specimens), primarily covering dorsal surface of base of Finger I, narrowing distally, extending to mid-proximal phalange on inner dorsal side; nuptial pad on Finger II small, oval, positioned on base of digit on inner dorsal side (extending to mid-proximal phalange on some individuals); external vocal sac indistinct; internal vocal slit present on floor of mouth near rear of mandible, one on each side; forearms enlarged relative to upper forelimbs. Females: mature ova without pigment; nuptial pads, vocal sac, vocal slits, and enlarged forearms, all absent. Morphological comparison. Megophrys major s.s. (adult males, N =12, adult females, N =3) differs from M. monticola and M. zhangi by its larger adult body size, male SVL 71.6–87.5 mm, female SVL 85.6–98.2 mm (vs. male SVL 38.2–49.5 mm, N =17, female SVL 40.5–56.1 mm, N =6; male SVL 32.5–37.2 mm, N =3, respectively); differs from M. robusta and M. medogensis by presence of distinct continuous whitish-cream upper lip stripe (vs. absent), by basal webbing present (vs. absent); differs from M. mangshanensis by basal webbing between toes present (vs. absent), lateral fringes on toes occasionally present (vs. absent), larger adult body size, male SVL 71.6–87.4 mm, female SVL 85.6–98.2 mm (vs. male SVL 62.5 mm, N=1, female SVL 73 mm, N=1). For comparisons with subsequent species covered in this study, refer to relevant morphological comparison sections for those species. Systematic position. OTU 5 (“ M. cf. major 3”) in the molecular analyses of this study and Mahony et al. (2017) represents Megophrys major s.s. The systematic position of M. major s.s. within the MMC clade is not fully resolved (Figures 2–5; Appendix I, Table 3; Appendix II, Figures 1, 2 & 5: as M. cf. major 3/OTU 5). It was consistently found to be the sister taxon to a new species (OTU 7/“ M. cf. major 1”) described below, but it is not yet clear which species these sister taxa are most closely related to within the MMC. Uncorrected p -distance for the 16S rRNA gene between these sister taxa was 6.4–6.5% (Appendix I, Table 6). Etymology. The original species epithet “ gigas ” is derived from the Greek word meaning “giant”, later replaced with “ major ”, Latin for “large”. Suggested common name: Jerdon’s White-lipped Horned Frog. Distribution. This species was considered widespread from Northeast India to southeastern Vietnam but M. major sensu lato (s.l.) is now known to represent a species complex (Chen et al. 2017; Mahony et al. 2017; this study). Only the following localities (between 450 and 1255 m asl.) represent the verified distribution of M. major s.s. (Figure 8A): Sohra [=Cherrapunji], in the East Khasi Hills district, Meghalaya state; Khumhzi, in Luangmai district and Khangchiuluanh, in Tamenglong district, Manipur state; and Tseminyu, in Kohima district, Nagaland. The locality “Darjeeling” associated with Jerdon’s BMNH series of specimens is considered erroneous (see Remarks for details). The eastern distribution limit for M. major s.s. is currently unclear, but it might be expected to extend into adjacent areas of western Myanmar. A report of this species from Tura Peak, West Garo Hills district in Meghalaya state was provided by Sangma and Saikia (2015). The authors provided no diagnostic characters for their specimens so the taxonomic identity of animals from this locality warrants careful investigation considering our study identifies the presence of three MMC species in Meghalaya state. Habitat and natural history. There have been many observations and notes published regarding M. major s.l., primarily from Southeast Asian countries (e.g., Bain & Nguyen 2004; Ohler et al. 2000; Stuart 1999, 2005), which are likely to represent different species. Some old reports of M. major tadpoles (e.g., Annandale 1912) from Northeast India are not considered reliable, since most tadpoles were identified by association of adult frogs present in the collection stream, not accounting for the fact that in our experience most streams in Northeast India are inhabited by two or more Megophrys species. Based on our observations of recently collected specimens, a female (CES 18901) collected towards the end of May contained mature ova, and males were found calling during May (CES 18902, CES 18903) and October (SDBDU 2007.229, SDBDU 2007.230), indicating that the breeding season for this species probably extends from the pre-monsoon (May) until post monsoon (October). SDBDU 2007.229 and SDBDU 2007.230 were found calling amongst fallen and dead reeds on the banks of the Terunhü River in Nagaland at around 18:30 h (just after dusk). The Terunhü River at the collection locality is a moderately fast flowing stream passing through a disturbed secondary forest patch. CES 18902 was found calling during the daytime perched ~ 2.3 m above ground level on a leaf of a wild cardamom on the bank of the Mpeih bak duithuak (stream). CES 18903 was found calling from a hole in the retaining wall where a clear water stream exits an underground diversion. The gravid female (CES 18901) was collected nearby an unnamed stream. No further individuals could be located along that stream during the subsequent two nights of searching, indicating that breeding activity had not yet reached a peak (or was experiencing a lull). Remarks. On Xenophrys gigas Jerdon, 1870: The original type series of Xenophrys gigas consisted of five syntype specimens in Jerdon’s personal collection collected from “ Sikim [sic] and the Khasi Hills” (Jerdon 1870). Soon after the description of Jerdon’s X. gigas, Anderson (1871b) stated “I have a specimen of Dr. Jerdon’s X. gigas beside me [in the Indian Museum (now ZSIC)], but I find that in no way differs from the Darjeeling large specimens, which are the adults of this species [Xenophrys monticola Günther, 1864]”. With that statement, X. gigas was regarded to represent a junior synonym of X. monticola for the subsequent four decades. Sclater (1892b:33) listed all Indian Museum amphibian specimens by numbers, localities and collectors, which included seven specimens of “ X. monticola ” sensu Anderson (1871b) deposited by Jerdon, six from “Darjeeling” (ZSIC 9673–9677, ZSIC 9721) and one from “Khasia Hills” (ZSIC 9679). We know now (discussed below) that the large species from Darjeeling that Anderson (1871b) was referring Jerdon’s X. gigas specimen to is M. robusta. However, it is not clear whether that particular specimen was one of Jerdon’s five syntype specimens of X. gigas. Deuti et al. (2017) referred five of the aforementioned “Darjeeling” specimens (ZSIC 9673–9677), and three others not mentioned by Sclater (1892b; ZSIC 9722–9724) to Megophrys sanu comb. nov. (as juveniles). The taxonomic identities of ZSIC 9721 and ZSIC 9679 require clarification. The whereabouts of the original type series of Xenophrys gigas, and which specimens actually belong to the original type series of this species, is an imbroglio. Chanda et al. (2000) referred four specimens (ZSIC 9670, ZSIC 9681, ZSIC 10777, ZSIC 10779) as types of X. gigas Jerdon, from “Darjeeling”, West Bengal, India. ZSIC also have an additional two specimens from Darjeeling that they regard to be the types of X. gigas (ZSIC 9650 and ZSIC 9668) (SM per. obs. during a visit to the ZSIC in 2010). Several problems exist with the recognition of these six ZSIC specimens as the types for this species: 1) all of these specimens were collected by J. Gammie (Sclater 1892b:33 [as Leptobrachium monticola]); Jerdon (1870) clearly stated that he “obtained” the type specimens from the type localities—Jerdon acknowledged throughout this paper when specimens were sent to him by other collectors, so it may be interpreted that he (or his party) were the collectors of the specimens; 2) ZSIC 9650 (SVL 51.1 mm) is a gravid adult female M. monticola (SM per. obs.). ZSIC 9668 was not examined in detail but is either an adult M. monticola or a juvenile M. robusta ––both specimens do not fit Jerdon’s original description by being &l, Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on pages 39-45, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/2610202, {"references":["Jerdon, T. C. (1870) Notes on Indian herpetology. part 2. Proceedings of the Asiatic Society of Bengal, March 1870, 66 - 85.","Boulenger, G. A. (1908) A revision of the oriental pelobatid batrachians (genus Megalophrys). Proceedings of the Zoological Society of London, 1908, 407 - 430, pls. XXII-XXV, fig. 71. https: // doi. org / 10.1111 / j. 1096 - 3642.1908. tb 01852. x","Mahony, S., Teeling, E. C. & Biju, S. D. (2013) Three new species of horned frogs, Megophrys (Amphibia: Megophryidae), from Northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region. Zootaxa, 3722 (2), 143 - 169. https: // doi. org / 10.11646 / zootaxa. 3722.2.2","Mahony, S., Foley, N. C., Biju, S. D. & Teeling, E. C. (2017) Evolutionary history of the Asian Horned Frogs (Megophryinae): integrative approaches to timetree dating in the absence of a fossil record. Molecular Biology and Evolution, 34 (3), 744 - 771 + S. I. 33 pp. https: // doi. org / 10.1093 / molbev / msw 267","Chen, J. - M., Zhou, W. - W., Poyarkov Junior, N. A. Stuart, B. L., Brown, R. M., Lathrop, A., Wang, Y. - Y., Yuan, Z. - Y., Jiang, K., Hou, M., Chen, H. - M., Suwannapoom, C., Ngoc Nguyen, S., Van Duong, T., Papenfuss, T. J., Murphy, R. W., Zhang, Y. - P. & Che, J. (2017) A novel multilocus phylogenetic estimation reveals unrecognized diversity in Asian horned toads, genus Megophrys sensu lato (Anura: Megophryidae). Molecular Phylogenetics and Evolution, 106, 28 - 43. https: // doi. org / 10.1016 / j. ympev. 2016.09.004","Sangma, M. A. & Saikia, P. K. (2015) Checklist of amphibians inventoried from Tura Peak of West Garo Hills, Meghalaya, India. Journal of Global Biosciences, 4 (5), 2404 - 2424.","Bain, R. H. & Nguyen, Q. T. (2004) Herpetofaunal diversity of Ha Giang Province in northeastern Vietnam, with descriptions of two new species. Novitates, 3453, 1 - 42. https: // doi. org / 10.1206 / 0003 - 0082 (2004) 453 2.0. CO; 2","Ohler, A., Marquis, O., Swan, S. & Grosjean, S. (2000) Amphibian diversity of Hoang Lien Nature Reserve (Lao Cai Province, northern Vietnam) with description of two new species. Herpetozoa, 13 (1 - 2), 71 - 87.","Stuart, B. L. (1999) Amphibians and reptiles. In: Duckworth, J. W., Salter, R. E. & Khounboline, K. (Eds.), Wildlife in Lao PDR. 1999 Status Report. IUCN-The World Conservation Union / Wildlife Conservation Society / Centre for Protected Areas and Watershed Management, Vientiane, pp. 43 - 67.","Stuart, B. L. (2005) New frog records from Laos. Herpetological Review, 36 (4), 473 - 479.","Annandale, N. (1912) Zoological results of the Abor Expedition, 1911 - 1912. I. Batrachia. Records of the Indian Museum, 8 (1), 7 - 36, pls. II-IV. https: // doi. org / 10.5962 / bhl. part. 1186","Anderson, J. (1871 b) On some Indian reptiles. Proceedings of the Zoological Society of London, 1871, 149 - 211.","Gunther, A. (1864) The Reptiles of British India. The Ray Society, London, 452 + XXVII pp., 26 pls.","Sclater, W. L. (1892 b) List of the Batrachia in the Indian Museum. Taylor and Francis, London, VIII + 43 pp.","Deuti, K., Grosjean, S., Nicolas, V., Vasudevan, K. & Ohler, A. (2017) Nomenclatural puzzle in early Xenophrys nomina (Megophryidae, Amphibia) solved with description of two new species from India (Darjeeling hills and Sikkim). Alytes, 34, 20 - 48.","Chanda, S. K., Das, I & Dubois, A. (2000 \" 1999 \") Catalogue of amphibian types in the collection of Zoological Survey of India. Hamadryad, 25 (2), 100 - 128.","Sclater, W. L. (1892 a) On some specimens of frogs in the Indian Museum, Calcutta with description of several new species. Proceedings of the Zoological Society of London, 1892, 341 - 348, pl. XXIV.","Blyth, E. (1855 [\" 1854 \"]) Notices and descriptions of various reptiles, new or little known. Journal of the Asiatic Society of Bengal, 23, 287 - 302.","Gunther, A. (1875) Second report on collections of Indian reptiles obtained by the British Museum. Proceedings of Zoological Society, 1875 (1), 224 - 234.","Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Caudata in the Collections of the British Museum. (Natural History). Second Edition. Taylor and Francis, London, XVI + 503 pp., XXX pls.","Gunther, A. (1876) Third report on collections of Indian reptiles obtained by the British Museum. Proceedings of Zoological Society, 1875 (4), 567 - 577.","Biju, S. D., Senevirathne, G., Garg, S., Mahony, S., Kamei, R. G., Thomas, A., Shouche, Y., Raxworthy, C. J., Meegaskumbura, M. & van Bocxlaer, I. (2016) Frankixalus, a new rhacophorid genus of tree hole breeding frogs with oophagous tadpoles. PLOS ONE, 11 (1), e 0145727. https: // doi. org / 10.1371 / journal. pone. 0145727","Gunther, A. (1860) Contributions to a knowledge of the reptiles of the Himalaya mountains. I. Descriptions of the new species. II. List of Himalayan reptiles, with remarks on their horizontal distribution. Proceedings of the Zoological Society of London, 1860, 148 - 175.","Daniel, J. C. (1962) Notes on some amphibians of the Darjeeling area, West Bengal. Journal of the Bombay Natural History Society, 59 (2), 666 - 668.","Subba, B., Aravind, N. A. & Ravikanth, G. (2016) Amphibians of the Sikkim Himalaya, India: an annotated checklist. Check List, 13 (1), 1 - 14."]}

Megophrys (Xenophrys) major s.s. Boulenger, 1908 (Figures 14 & 15; Table 1) Xenophrys gigas Jerdon 1870:85 (partim: Khasi Hills). In: Notes on Indian herpetology. Proceedings of the Asiatic Society of Bengal, March, 1870: 66���85. Megalophrys major Boulenger 1908:410, 416, Pl. xxiii. In: A revision of the oriental pelobatid batrachians (genus Megalophrys). Proceedings of the Zoological Society of London, 1908: 407���430 + Pl. xxii���xxv + fig. 71., Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on page 39, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/2610202, {"references":["Boulenger, G. A. (1908) A revision of the oriental pelobatid batrachians (genus Megalophrys). Proceedings of the Zoological Society of London, 1908, 407 - 430, pls. XXII-XXV, fig. 71. https: // doi. org / 10.1111 / j. 1096 - 3642.1908. tb 01852. x","Jerdon, T. C. (1870) Notes on Indian herpetology. part 2. Proceedings of the Asiatic Society of Bengal, March 1870, 66 - 85."]}

Megophrys (Xenophrys) oreocrypta sp. nov. (Figures 18 & 19; Table 1) Holotype. One adult female (BNHS 6045 [field no. SDBDU 2009.1108]: Figures 18, & 19A & C), from Tura Peak Reserve Forest (25��30'29.4"N, 90��13'44.82"E, 640 m asl.), near Tura town, West Garo Hills district, Meghalaya state, Northeast India, collected by members of the Systematics Lab, University of Delhi on 20 June 2009. Paratypes. Four unsexed juveniles (BNHS 6046 ��� BNHS 6049 [field nos. SDBDU 2009.1104 ��� 1107]: Figure 19B, D, E & F), collected along with the holotype. Referred specimen. One unsexed juvenile (SDBDU 2008.1400; not measured), from Tura Peak Reserve Forest (25��30'40"N, 90��13'50"E, 510 m asl.), near Tura town, West Garo Hills district, Meghalaya state, Northeast India, collected by RGK on 13 May 2008; one juvenile (FMNH 74154 [previously ZSIC 18492]), from ���Tura, Garo Hills, Assam ��� (now West Garo Hills district, Meghalaya state), collected by S.W. Kemp, date unknown. Holotype description (measurements in mm). Mature female (SVL 94.9) (Figures 18, & 19A & C). Head moderately large, slightly wider than long (HW 35.9, HL 35.7, IFE 16.2, IBE 25.6); snout bluntly pointed in dorsal view, obtusely protruding beyond mandible in lateral view, without rostral appendage (Figure 18C); loreal region acute, concave; canthus rostralis angular; dorsal surface of snout concave; eye diameter almost twice as long as maximum diameter of visible portion of tympanum, and shorter than length of snout (EL 9.0, TYD 4.7, SL 11.8); eye���tympanum distance (TYE 8.2) longer than diameter of visible portion of tympanum; tympanum oval-shaped, slightly oblique (Figure 18C), with upper ~15% concealed by supratympanic ridge; pupil in life vertically elliptical; nostril positioned laterally, closer to eye than to snout (EN 6.0, NS 6.5); internarial distance greater than eyelid width, and slightly less than narrowest point between upper eyelids (IN 9.3, UEW 8.4, IUE 9.6); pineal ocellus not visible externally; vomerine ridges medium sized, ovoid, moderately raised, positioned between to slightly posterior to choanae, equidistant from each other and choanae; vomerine teeth small; maxillary teeth present; tongue moderately large, medial lingual process absent, posterior edge not observed due to fixation of jaw and in interest of preventing potential damage to jaws by forcing open mouth wide enough for examination. Forelimbs moderately long, thin (Figures 18A & B, & 19A & C), forearm not enlarged relative to upper forelimb, and shorter than hand length (FAL 21.5, HAL 23.3); fingers moderately long, narrow, without lateral fringes (Figure 18D), finger length formula IV Colouration: In preservative (Figure 18): Dorsal and lateral surfaces of body, and dorsal surface of head primarily mid brown; light-edged, dark brown incomplete triangular marking between eyes; faint dark brown blotch on centre of dorsum; tubercles on flanks primarily creamish-white; lateral surfaces of head below supratympanic ridges and canthus rostralis dark brown with cream-white stripe on upper lip, extending from anterior to nostril to rear of jaw; dorsal and lateral surfaces of forelimbs and hindlimbs primarily brown; two or three dark brown blotches on dorsolateral surface of forearms; dorsal surface of fingers with dark brown blotches; dorsal surfaces of hindlimbs with faint transverse crossbars; throat pale brown with brown and creamish-white spots and blotches along edge of gular region, light-edged wide light brown stripe extending from posterior edge of mandible onto base of forearms; chest light brown; abdomen, ventral surfaces of forelimbs and thighs plain creamish-white, ventral surface of shanks grey; dark brown longitudinal stripe ventrolaterally on both sides of abdomen; area surrounding vent and posterior surfaces of thighs dark brown; ventral surfaces of tarsi and feet dark greyish-brown; hands ventrally pale greyish-brown; pectoral and femoral glands creamish-white. In life (Figure 19A & C): Dorsal surfaces of head, body and limbs orangish-brown; upper lip stripe pale greyish-brown, darker anteriorly, fading posteriorly; posterior flanks and groin region with yellow hue; ventral surface of throat and chest primarily pale grey with dark orange speckling, speckling more dense on lateral stripes of throat, and ventrolaterally on anterior abdomen; abdomen and ventral surfaces of thighs white. Variation. See Table 1 for morphometric variation within the examined specimen series, consisting of an adult female, and five juveniles. All juveniles have head width less than head length; FMNH 74154 has finger length formula IVSecondary sexual characters. Males currently unknown. Morphological comparison. Megophrys oreocrypta sp. nov. (adult female, N =1) differs from M. monticola by its larger adult female body size, SVL 94.9 mm (vs. female SVL 40.5���56.1 mm, N =6); differs from M. robusta, M. medogensis and M. zhangi by absence of distinct dark brown vertical bar extending from lower orbital edge to edge of upper lip (vs. present) and presence of a distinctly lighter continuous upper lip stripe (vs. upper lip stripe absent). From other members of M. major species complex, it can be distinguished from Megophrys flavipunctata sp. nov. and M. mangshanensis by its larger adult female body size, SVL 94.9 mm (vs. female SVL 68.0��� 74.6 mm, N =3; female SVL 73.0 mm, N =1, respectively), upper lip stripe extends anteriorly beyond nostril, N =4 [includes subadults] (vs. stripe terminates at, or posterior to nostril); differs from M. major s.s. by larger tympanum diameter/ eye diameter ratio, TYD/EL 52.2% (vs. female TYD/EL 40.0���41.2%, N =3), upper lip stripe extends anteriorly beyond nostril, N =4 [includes subadults] (vs. stripe terminates at, or posterior to nostril on all examined specimens, N= 15), pair of distinctly enlarged glands adjacent to cloaca absent, N =2 (vs. present, N =15). Systematic position. This taxon represents M. cf. major 1 in the molecular analyses (discussed as OTU 7), and in Mahony et al. (2017). The systematic position of Megophrys oreocrypta sp. nov. within the MMC clade is not fully resolved. It has consistently been found to be a member of the western clade of the MMC, and the sister taxon to M. major s.s. across all multigene analyses except the ML analysis of Dataset I (Appendix II, Figure 6), however, the relationship of this sister taxa clade with other members of the MMC is not clear when molecular data for the MMSG was subjected to a variety of phylogenetic techniques (Figures 2���5; Appendix I, Table 3; Appendix II, Figures 1, 2, 5 & 6). Refer to Appendix I, Table 6 for uncorrected p -distances for the 16S rRNA gene between Megophrys oreocrypta sp. nov. and other MMSG species. Etymology. The specific epithet ��� oreocrypta ��� is an adjective derived from the Latinised Greek words ��ros meaning ���mountain��� and kryptos meaning ���hidden��� or ���secret���, in reference to the difficulty of finding adults of this species despite making several field trips to the Tura Peak Reserve Forest during 2008, 2009 and 2011. Suggested common name: Garo White-lipped Horned Frog. Sangma and Saikia (2015) provide the local Garo (language) name ���Diplok Nakma��� for this species. Distribution. Megophrys oreocrypta sp. nov. is currently known only from the southern slopes of the Tura Peak Reserve Forest (510���640 m asl.), in the West Garo Hills district of Meghalaya state (Figure 8A). Sangma and Saikia (2015) collected this species from slightly lower elevation on Tura Peak (437 m asl.) at Rengsangrap waterfall. It is likely that this species will be found throughout the adjoining Nokrek National Park and Biosphere Reserve, however its eastern distribution limits and altitudinal range are yet to be determined. Habitat and natural history. The habitat at the collection locality for this species consisted primarily of tropical lowland forest. The adult female holotype was collected after dusk, from the banks of one of the major streams flowing west towards Tura town. Juveniles were all collected after sunset on forest trails within ~ 20 m from the nearest large stream. The biology of this species remains unknown; no adult males have been heard calling from streams that were surveyed and the ovaries of the holotype are empty. Sangma and Saikia (2015) reported observing two (or five, according to their table 1) individuals of this species (as Xenophrys glandulosa) at the type locality, from among leaf litter and under moist rocks in the forest. Remarks. A report of M. robusta by Dutta (1997) from the Garo Hills is presumably based on Megophrys oreocrypta sp. nov. ( see ���Remarks��� for M. robusta). Smith (1929) reported M. major from Tura possibly based on the specimen collected by W. Kemp (FMNH 74154 [originally ZSIC 18492]), which we herein verify to represent Megophrys oreocrypta sp. nov. Sangma and Saikia (2015) provided a photograph (plate 4a) of a preserved specimen collected from the type locality, Tura Peak, which they referred to as Megophrys glandulosa. This specimen appears to represent Megophrys oreocrypta sp. nov. based on the presence of a white upper lip stripe that clearly extends anteriorly beyond the nostril., Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on pages 50-54, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/2610202, {"references":["Mahony, S. (2008) Redescription and generic reallocation of Rana mawphlangensis Pillai & Chanda, 1977 (Amphibia: Ranidae). Hamadryad, 33 (1), 1 - 12.","Mahony, S., Foley, N. C., Biju, S. D. & Teeling, E. C. (2017) Evolutionary history of the Asian Horned Frogs (Megophryinae): integrative approaches to timetree dating in the absence of a fossil record. Molecular Biology and Evolution, 34 (3), 744 - 771 + S. I. 33 pp. https: // doi. org / 10.1093 / molbev / msw 267","Sangma, M. A. & Saikia, P. K. (2015) Checklist of amphibians inventoried from Tura Peak of West Garo Hills, Meghalaya, India. Journal of Global Biosciences, 4 (5), 2404 - 2424.","Dutta, S. K. (1997) Amphibians of India and Sri Lanka (Checklist and Bibliography). Odyssey Publishing House, Bubaneswar, XIII + 342 + XXII pp.","Smith, M. A. (1929) On a collection of amphibians and reptiles from the upper reaches of the Brahmaputra. Records of the Indian Museum, 31, 77 - 80."]}

Megophrys (Xenophrys) robusta Boulenger, 1908 (Figures 11 & 12; Table 1) [?] Xenophrys gigas Jerdon 1870:85 (partim: ��� Sikim ���). In: Notes on Indian herpetology. Proceedings of the Asiatic Society of Bengal, March, 1870: 66���85. Megalophrys robusta Boulenger 1908:418, pl. xxiv. In: A revision of the oriental pelobatid batrachians (genus Megalophrys). Proceedings of the Zoological Society of London, 1908: 407���430 + pl. xxii���xxv + fig. 71. Lectotype (by present designation). Adult female (BMNH 1947.2.25.19 [RR 1908.4.8.8]: Figure 11), from ���Darjeeling��� (~ 27��03'0"N, 88��16'0"E), West Bengal, India, collector J. Gammie, presented by the Indian Museum, collection date unknown. Paralectotypes (by implication). Two adult females (ZSIC 9681, ZSIC 10777), details are the same as for the lectotype. Examined specimens. BMNH 1947.2.25.19 (lectotype: Figure 11); ZSIC 9681, ZSIC 10777 (paralectotypes); three adult males (SDBDU 2011.1057; SDBDU 2011.1062: Figure 12A; SDBDU 2011.1064: Figure 12F), and one adult female (SDBDU 2011.1063: Figure 12C & F), from near Latpanchar town (26��54'34.2"N, 88��23'54.54"E, 1135 m asl.), Kurseong sub-division, Darjeeling district, West Bengal, Northeast India, collected by the Systematics Lab members on 0 5 and 0 6 June 2011; a subadult male (SDBDU 2011.415: Figure 12E), and an adult male (SDBDU 2011.416: Figure 12E), from Stream 1 (26��54'34.7"N, 88��23'52.7"E, 1030 m asl.), Mahananda Wildlife Sanctuary, Latpanchar township, Kurseong sub-division, Darjeeling district, West Bengal, Northeast India, collected by SDB and RGK on 19 and 20 May 2011; two adult males (SDBDU 2009.1270; SDBDU 2009.1284: Figure 12B), and one juvenile (SDBDU 2009.1245: Figure 12D), from Sessa village (27��6'4.02"N, 92��31'38.52"E, 1110 m asl.), West Kameng district, Arunachal Pradesh state, Northeast India, collected by the Systematics Lab members on 07���09 August 2009. Lectotype description (measurements in mm). Mature female (SVL 102.0) (Figure 11). Head large, wider than long (HW 42.0, HL 40.9, IFE 16.9, IBE 30.1); snout broadly pointed in dorsal view, obtusely protruding beyond mandible in lateral view, without rostral appendage (Figure 11C); loreal region acute, concave, with well developed canthus rostralis; dorsal surface of snout concave; eye diameter larger than maximum tympanum diameter, and shorter than snout (EL 11.6, TYD 7.7, SL 13.2); eye���tympanum distance (TYE 8.2) slightly longer than tympanum diameter; tympanum distinctly oval, obliquely orientated (Figure 11C), upper ~40% concealed by supratympanic ridge; pupil indistinct; nostril positioned laterally, circular with raised posterior rim, closer to eye than to snout (EN 5.3, SN 7.4); internarial distance equal to narrowest point between upper eyelids, and greater than upper eyelid width (IN 11.9, IUE 12.0, UEW 10.0); pineal ocellus absent; vomerine ridges well developed, positioned between choanae, closer to choanae than to each other; vomerine teeth absent; maxillary teeth present; tongue large, rounded posteriorly (possibly artefact of preservation), medial lingual process absent. Forelimbs moderately long, thin (Figure 11A & B), forearm slightly enlarged relative to upper forelimb, slightly shorter than hand length (FAL 25.4, HAL 26.3); fingers long, narrow (Figure 11D), finger length formula IV Colouration: In preservative (Figure 11): Flanks, and dorsal surface of head, body, and limbs primarily plain brown; large flank tubercles lighter brown, each bordered below by small dark brown blotch; faint solid dark brown triangular marking on dorsal surface of head; wide, slightly oblique dark brown bar below each eye; gular region and chest pale brown with some dark speckling; ventral surfaces of thighs and shanks primarily light with faint mottling; outer tarsi with continuous dark brown blotch extending from base of foot to base of shank along outer edge; surfaces adjacent to cloaca dark brown; forelimbs brown above with two or three dark brown transverse stripes and blotches on forearms; dark brown patch extends from base of hand to elbow on ventral surface; ventral surfaces of hands and feet faded greyish-brown; pectoral and femoral glands white. In life: Not originally documented for lectotype (refer to Figure 12 for other examples). Variation. Refer to Table 1 for morphometric variation within the type series and referred specimens, consisting of seven adult males, four adult females, a subadult, and a juvenile. The paralectotypes and referred specimens resemble the lectotype for most morphological characters with the following exceptions: Head width/ head length ratios differ between Darjeeling and Arunachal Pradesh specimens���all Darjeeling specimens have HW&Zcaron;HL (this study, N =9; Deuti & Kumar 2008, N =5), Arunachal Pradesh specimens have HL&Zcaron;HW (this study, N =3); two specimens (SDBDU 2009.1270, SDBDU 2009.1245) have relative finger length formula IV=I=II (���, ���>--M. robusta by Ohler et al. (2002)] (Figures 11C, & 12A, C & E); outer edge of gular region typically brown, with eight light patches. Secondary sexual characters. Males: nuptial pads present, weakly raised, covered with black microasperities, covering almost entire dorsal surface of Finger I; nuptial pad on Finger II small, oval, positioned on base of digit on inner dorsal side (extending on to base of proximal phalange on some individuals); external vocal sac usually indistinct, though skin is loose on some individuals indicating large sub-gular sac that extends onto chest; large internal vocal slit is present on floor of mouth near rear of mandible on each side; forearms enlarged relative to upper forelimbs. Females: ova not pigmented; nuptial pads, vocal sac, vocal slits, all absent. Morphological comparison. Megophrys robusta (adult males N =6, adult females N =4) differs from M. monticola and M. zhangi by its larger adult body size, male SVL 73.5���83.1 mm, female SVL 81.3���108.3 mm (vs. male SVL 38.2���49.5 mm, N =17, female SVL 40.5���56.1 mm, N =6; male SVL 32.5���37.2 mm, N =3, respectively); differs from M. mangshanensis by absence of distinct white upper lip stripe (vs. present). For comparisons with subsequent species covered in this study, refer to relevant morphological comparison sections for those species. Systematic position. Megophrys robusta was found to be the sister taxon to the MMC clade, and was the second most ancestrally derived lineage in the MMSG after M. monticola (Mahony et al. 2017; Figures 2 & 3; Appendix II, Figures 2, 3A & 4B). The inclusion of additional sequences from Chen et al. (2017) in ML analyses demonstrates that M. robusta is the sister taxon to the morphologically similar M. medogensis, however, the systematic position of this clade was not strongly supported (Figure 4; Appendix II, Figures 5 & 6). Refer to Appendix I, Table 6 for uncorrected p -distances for the 16S rRNA gene between M. robusta and other MMSG species. Etymology. The specific epithet ��� robusta ��� is Latin for ���hard��� or ���strong���, presumably a reference to the heavy set habitus of this species. Suggested common name: Robust Horned Frog. Distribution. Specimens examined in this study demonstrate that M. robusta ranges at least from the Darjeeling Hills, northern West Bengal state, east through Bhutan to at least the West Kameng district of western Arunachal Pradesh state, in India (Figure 8A). Specimens examined in this study were collected at elevations from 1030 to 1135 m asl. Daniel (1962) reported two specimens (as M. major) collected from Darjeeling itself (~ 2200 m asl.) and from nearby Darjeeling (~ 1675 m asl.), Darjeeling Sadar division, Darjeeling district, West Bengal. Rai and Anders (2002) provided a number of localities in Ilam, Pachthar and Taplejung provinces in eastern-most Nepal (bordering Darjeeling and Sikkim) from 1600 to 2400 m asl., indicating that the Arun River forms the western boundary for the species. They provided a description of M. robusta and figures, which corresponded well with the species as defined here. This species is also found in neighbouring Sikkim state, in Northeast India (Subba et al. 2016; Deuti et al. 2017), and may extend north along river valleys into bordering southern Tibet, China. Refer to the Remarks section for discussions on additional localities published elsewhere. Habitat and natural history. All but one of the specimens collected during this study were collected after dusk from the banks of moderate sized (3���10 m wide), typically shallow rocky hill streams surrounded by at least patches of mature forest, and dense vegetation (Figure 9A). In Darjeeling district during late May to early June, calls provisionally attributed to this species (a loud typical Megophrys (Xenophrys) call) were occasionally heard from streams during both day and night surveys. These calls consisted of a short burst of notes followed by a long silence. At Sessa, similar calls were heard during night surveys in early August, and were used on two occasions to locate individuals. However, in both instances the suspected calling males were found within 1���2 m of a similar sized, and considerably more abundant sympatric MMC species (described below) on the banks of the Sessa River, so calls could not be confirmed to be from M. robusta. At both localities, adult male individuals were always widely dispersed along stream banks and were typically found perched on top of large rocks in the open. A female (SDBDU 2011.1063, SVL 81.3 mm: Figure 12C & F) was collected from a roadside embankment amongst dense vegetation ~ 50 m from the nearest stream. Its ovaries contained small, pigmentless ova. Daniel (1962) reported that a large female (SVL 101 mm) collected in June contained enlarged ova (diameter 2 mm), however, he stated a female specimen (SVL 80 mm) ���collected in September had the ovaries dormant���. Though the latter female may have been sexually immature, the available evidence suggests that the breeding season for this species extends at least from mid-May until August. A male specimen (SDBDU 2011.415, SVL 69.1 mm: Figure 12E) collected in late May had distinctly enlarged testes, but had not yet developed nuptial pads and the internal vocal slits were small indicating that this was not yet sexually mature. SDBDU 2009.1284 (SVL 73.5 mm [Figure 12B]) was the smallest sexually mature male. Rai and Anders (2002) provided a number of additional observations that are of interest, however, their description of the call (as ���a sharp whistling call, sounding like ��� khui-khui ���) does not correspond with the authors��� experience, where calls were distinctly of a lower frequency sound more similar to the ���caws��� of a crow. Remarks. Boulenger (1908) described this species based on a syntype series of five specimens collected by Dr. J. Gammie from Darjeeling (four in the Indian Museum [now ZSIC], one in the BMNH [now NHMUK]���the figured specimen). He provided measurements of two ZSIC specimens, a female (SVL 114 mm), and a ���halfgrown��� individual (SVL 54 mm). Dutta (1997) erroneously claimed (and followed by Bordoloi & Borah 2001) that M. robusta was a replacement name for X. gigas Jerdon, 1870 (now M. major). Dutta (1997) also only mentioned ZSIC as the location of type specimens. Chanda et al. (2000), apparently also following Dutta (1997), did not mention M. robusta in the list of type specimens in ZSIC, but instead assigned four specimens (ZSIC 9670, ZSIC 9681, ZSIC 10777, ZSIC 10779) as syntypes of Xenophrys gigas. ZSIC 9681 and ZSIC 10777 clearly represent M. robusta. ZSIC 9670 and ZSIC 10779 were not examined in this study. The ZSIC also recognised (according to a jar label) two additional specimens, ZSIC 9650 (an adult female M. monticola, SVL 51.1 mm) and ZSIC 9668 (not examined in detail but represents either an adult M. monticola or juvenile M. robusta) as ���types��� of X. gigas. Our study demonstrates that none of these ZSIC specimens are the types of X. gigas (see X. major Remarks section). Thus it remains unclear which four of these six specimens represent the Indian Museum (ZSIC) syntypes mentioned by Boulenger (1908) for M. robusta. For this reason, only BMNH 1908.4.8.8 can currently be verified as a syntype of M. robusta and is therefore designated above as the lectotype of this taxon, thus providing nomenclatural stability by restricting the name to a verified original syntype. Megophrys robusta has been reported from several localities further east in Arunachal Pradesh state, e.g., Lower Subansiri district (Bordoloi et al. 2000; Bordoloi & Borah 2001) and the Upper Siang district and Changlang districts (Pawar & Birand 2001). Pawar and Birand (2001) did not provide a description of the species they observed and did not appear to have collected voucher specimens, so these localities should be considered anecdotal pending collection of verifiable specimens from the Upper Siang and Changlang districts. Several of the morphological characters described by Bordoloi and Borah (2001) indicated that they may have misidentified an MMC species, e.g., ���belly with faint darker spots���. They however did not provide further defining and useful taxonomic characters to help identify which species they collected. Therefore, the identities of Bordoloi and Borah���s (2001) specimens require verification before Lower Subansiri district can be included in the verified distribution of M. robusta. Dutta (1997) reported M. robusta from the Garo Hills in Meghalaya state, presumably misidentifying the large unnamed MMC species (formally described below), so this locality also requires verification based on the reexamination of voucher specimens. Sangma and Saikia (2015) provided an account and figure of a specimen they referred to as M. robusta from Tura Peak, West Garo Hills district in Meghalaya state. This specimen clearly possesses a whitish upper lip stripe (absent on M. robusta), demonstrating that it is one of the three MMC sp, Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on pages 30-37, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/2610202, {"references":["Boulenger, G. A. (1908) A revision of the oriental pelobatid batrachians (genus Megalophrys). Proceedings of the Zoological Society of London, 1908, 407 - 430, pls. XXII-XXV, fig. 71. https: // doi. org / 10.1111 / j. 1096 - 3642.1908. tb 01852. x","Jerdon, T. C. (1870) Notes on Indian herpetology. part 2. Proceedings of the Asiatic Society of Bengal, March 1870, 66 - 85.","Deuti, K & Kumar, A. (2008) Rediscovery of Robust pelobatid frog, Xenophrys robusta, after fifty years with three colour morphs. Zoological Survey of India, 108 (4), 119 - 120.","Ohler, A., Swan, S. R. & Daltry, J. C. (2002) A recent survey of the amphibian fauna of the Cardamom Mountains, southwest Cambodia with descriptions of three new species. The Raffles Bulletin of Zoology, 50 (2), 465 - 481.","Mahony, S., Foley, N. C., Biju, S. D. & Teeling, E. C. (2017) Evolutionary history of the Asian Horned Frogs (Megophryinae): integrative approaches to timetree dating in the absence of a fossil record. Molecular Biology and Evolution, 34 (3), 744 - 771 + S. I. 33 pp. https: // doi. org / 10.1093 / molbev / msw 267","Chen, J. - M., Zhou, W. - W., Poyarkov Junior, N. A. Stuart, B. L., Brown, R. M., Lathrop, A., Wang, Y. - Y., Yuan, Z. - Y., Jiang, K., Hou, M., Chen, H. - M., Suwannapoom, C., Ngoc Nguyen, S., Van Duong, T., Papenfuss, T. J., Murphy, R. W., Zhang, Y. - P. & Che, J. (2017) A novel multilocus phylogenetic estimation reveals unrecognized diversity in Asian horned toads, genus Megophrys sensu lato (Anura: Megophryidae). Molecular Phylogenetics and Evolution, 106, 28 - 43. https: // doi. org / 10.1016 / j. ympev. 2016.09.004","Daniel, J. C. (1962) Notes on some amphibians of the Darjeeling area, West Bengal. Journal of the Bombay Natural History Society, 59 (2), 666 - 668.","Rai, K. R. & Anders, C. (2002) Megophrys robusta (Boulenger, 1908). In: Schleich, H. H. & Kastle, W. (Eds.), Amphibians and Reptiles of Nepal. A. R. G. Gantner Verlag K. G., Ruggel, pp. 167 - 171.","Subba, B., Aravind, N. A. & Ravikanth, G. (2016) Amphibians of the Sikkim Himalaya, India: an annotated checklist. Check List, 13 (1), 1 - 14.","Deuti, K., Grosjean, S., Nicolas, V., Vasudevan, K. & Ohler, A. (2017) Nomenclatural puzzle in early Xenophrys nomina (Megophryidae, Amphibia) solved with description of two new species from India (Darjeeling hills and Sikkim). Alytes, 34, 20 - 48.","Dutta, S. K. (1997) Amphibians of India and Sri Lanka (Checklist and Bibliography). Odyssey Publishing House, Bubaneswar, XIII + 342 + XXII pp.","Bordoloi, S. & Borah, M. M. (2001) First record of a frog Megophrys robusta (Boulenger 1908) from Arunachal Pradesh, India. Cotton College Research Journal, 1, 191 - 193.","Chanda, S. K., Das, I & Dubois, A. (2000 \" 1999 \") Catalogue of amphibian types in the collection of Zoological Survey of India. Hamadryad, 25 (2), 100 - 128.","Bordoloi, S., Borah, M. M., Sharma, K. K. & Kalita, S. N. (2000) An overview of amphibian fauna of Arunachal Pradesh, India. Journal of the Assam Science Society, 41 (4), 363 - 371.","Pawar, S. & Birand, A. (2001) A Survey of Amphibians, Reptiles, and Birds of Northeast India. CERC Technical Report 6. Center for Ecological Research and Conservation, Mysore, 75 pp.","Sangma, M. A. & Saikia, P. K. (2015) Checklist of amphibians inventoried from Tura Peak of West Garo Hills, Meghalaya, India. Journal of Global Biosciences, 4 (5), 2404 - 2424.","Hora, S. L. (1923) Observations on the fauna of certain torrential streams in the Khasi Hills. Records of the Indian Museum, 25, 579 - 599.","Khan, M. M. H. (2008) Protected Areas of Bangladesh; A Guide to Wildlife. Nishorgo Programme, Dhaka, 305 pp.","Sen, N. & Mathew, R. (2008) Bibliographical notes on the amphibians of Northeast India. Records of the Zoological Survey of India Occasional Papers, 291, 1 - 58."]}

Megophrys (Xenophrys) periosa sp. nov. (Figures 22 & 23; Table 1) Holotype. Adult male (BNHS 6055 [field no. SDBDU 2009.793]: Figures 22, & 23A & H), from Pangin town (28°12'33.96"N, 94°59'10.02"E, 450 m asl.), East Siang district, Arunachal Pradesh state, Northeast India, collected by members of the Systematics Lab, University of Delhi on 26 July 2009. Paratypes. Four adult males (BNHS 6057 [field no. SDBDU 2009.794]: Figure 23B; BNHS 6058–6060 [field nos. SDBDU 2009.1170 – 1172]: Figure 23H), collected along with the holotype by members of the Systematics Lab, University of Delhi on 26–28 July 2009; four adult males (BNHS 6061 [field no. SDBDU 2009.1243]: Figure 23E; BNHS 6062 [field no. SDBDU 2009.1244], BNHS 6063–6064 [field nos. SDBDU 2009.1265 – 1266]: Figure 23G), and one adult female (BNHS 6056 [field no. SDBDU 2009.1285]: Figure 23F), from the Sessa River, nearby Sessa village (27°6'4.02"N, 92°31'38.52"E, 1110 m asl.), West Kameng district, Arunachal Pradesh state, Northeast India, collected by members of the Systematics Lab, University of Delhi on 07–09 August 2009. Referred specimens. Two adult males (SDBDU 2009.1189: Figure 23D; SDBDU 2009.1190: Figure 23C), from Rigo Village (28°9'34.56"N, 94°47'19.38"E, 260 m asl.), Along town, West Siang district, Arunachal Pradesh state, Northeast India, collected by Systematics Lab members on 30 July 2009; one adult male (SDBDU 2009.1267), from the Sessa River, nearby Sessa village (27°6'4.02"N, 92°31'38.52"E, 1110 m asl.), West Kameng district, Arunachal Pradesh state, Northeast India, collected by members of the Systematics Lab, University of Delhi on 07–09 August 2009. Provisionally referred specimens (see Remarks). One juvenile male (SDBDU 2009.132), and two juvenile females (SDBDU 2009.133 & SDBDU 2009.134), from Pangkava ju (24°40'21.6"N, 94°28'19.86"E, 820 m asl.), Kangpat Khullen, Kamjong sub-division, Ukhrul district, Manipur state, Northeast India, collected by RGK and SDB on 21 May 2009; adult female (CAS 232938 ––tissue only), from Hepu Stream (25°5'25.2"N, 96°24'13.2"E), Hepu village, Moe Nyin township, Myitkyina district, Kachin state, northern Myanmar, collected by G.O.U. Wogan, J.A. Wilkinson, J.V. Vindum, H. Win, T. Thin [“Additional collectors: K.S. Lwin, A.K. Shein and H. Tun ”] on 14 May 2003. Holotype description (measurements in mm). Mature male (SVL 93.8) (Figures 22, & 23A & H). Head moderately large, wider than long (HW 35.0, HL 33.6, IFE 15.5, IBE 25.3); snout bluntly pointed in dorsal view, obtusely protruding beyond mandible in lateral view, without rostral appendage (Figure 22C); loreal region acute, concave; canthus rostralis angular; dorsal surface of snout very slightly concave; eye diameter twice maximum diameter of visible portion of tympanum, and shorter than snout length (EL 10.7, TYD 5.4, SL 11.7); eyetympanum distance (TYE 8.2) less than twice diameter of visible portion of tympanum; tympanum oval-shaped, oblique (Figure 22C), with upper ~15% concealed by supratympanic ridge; pupil vertically elliptical; nostril positioned laterally, closer to eye than to snout (EN 5.2, NS 7.1); internarial distance greater than eyelid width, and equal to narrowest point between upper eyelids (IN 10.5, UEW 8.7, IUE 10.5); pineal ocellus not visible externally; vomerine ridge present, medium sized, ovoid, moderately raised, orientated acutely, positioned between to slightly posterior to choanae, equidistant from choanae and each other; vomerine teeth short; maxillary teeth present; tongue not observed due to fixation of jaw and in interest of preventing potential damage to jaws by forcing open mouth wide enough for examination. Forelimbs long, thin (Figures 22A & B, & 23A), forearm moderately enlarged relative to upper forelimb, and shorter than hand length (FAL 20.6––left side; right side deformed; HAL 24.0); fingers long, narrow, without lateral fringes (Figure 22D), finger length formula IV Skin of dorsal and lateral surfaces of head, body and limbs primarily smooth, sparsely covered with small, weak granules; tympanum with borders raised relative to surrounding region; outer edge of upper eyelids with a broad pointed bump; supratympanic ridges narrow anteriorly, gradually expanding posterior to tympanum to become moderately enlarged, glandular, extending from orbit, curving downward abruptly at posterior upper border of tympanum, terminating above forelimb insertions; flanks sparsely covered with small to medium sized unevenly scattered pustular tubercles; dorsolateral ridges thin, weakly developed, extending posteriorly from behind supratympanic ridges to ~90% distance to vent; parietoscapular-sacral ridges weakly developed, “>– Colouration: In preservative (Figure 22): Dorsal surfaces of body and head primarily greyish-brown; lightedged, darker brown triangular marking between eyes; parietoscapular-sacral and dorsolateral ridges narrowly bordered by dark brown; flanks grey, darker dorsally, tubercles on flanks primarily dark brown or black, many with small lighter tips; large dark brown blotch covers tympanic region, another extends from lower posterior border of orbits, additional dark brown bar extends vertically from lower orbital border to edge of upper lips; lateral surfaces of canthus rostralis black, front of snout dark brown; remaining surfaces of upper lips and area between dark blotches on lateral surfaces of head greyish-white; dorsal and lateral surfaces of forelimbs primarily light grey with large dark brown transverse blotches on forearms; dorsal surface of fingers with dark brown blotches; dorsal and lateral surfaces of hindlimbs brown with distinct dark brown transverse crossbars; throat, chest and anterior abdomen primarily brown with white-edged darker brown blotches and mottling; posterior half of abdomen and ventral surfaces of forelimbs and hindlimbs primarily creamish-white with dark brown blotches; ventral surfaces of tarsi dark brown; area surrounding vent and posterior surfaces of thighs dark brown; ventral surfaces of feet and hands greyish-brown; pectoral and femoral glands creamish-white. In life (Figure 23A): Dorsal surfaces of head, body and hindlimbs more olive-brown, distinctly paler on flanks and forelimbs; ventral surfaces generally lighter, with darker brown blotches distinct and revealing distinct wide brown longitudinal stripe on gular region; dark blotches on chest; ventral surfaces of limbs dark to light purple/mauve; distinct orange speckles present on dorsal and ventral surfaces, most noticeable on dorsal surfaces of toes and fingers and on flanks; iris very dark orange. Refer to Figures 22 and 23A for further details of colouration and markings for holotype (in life, and in preservation). Variation. Refer to Table 1 for morphometric variation within the type series and referred specimens, consisting of 12 adult males and one adult female. The paratypes and referred specimens generally resemble the holotype for most morphological characters with the following exceptions: Lateral fringes on toes typically absent, but some specimens have very narrow fringes on all toes; relative finger lengths vary between individuals, including IV75% of the trunk length (Figure 23 A–F); typically only upper border of tympanum is concealed by supratympanic ridge, though up to ~25% of tympanum appears to be concealed on some specimens; parietoscapular-sacral ridge configuration varies considerably amongst individuals, i.e., “>- (”, “>–– |”, “>- Secondary sexual characters. Males: nuptial pads present, weakly raised, covered with brown/black microasperities on freshly collected specimens, covering most of dorsal surface of Finger I; nuptial pad on Finger II medium sized, oval, positioned on base of digit on inner dorsal side, extending almost to base of distal phalange; external vocal sac indistinct; large internal vocal slits present on floor of mouth near rear of mandible, on each side; forearms enlarged relative to upper forelimbs. Female: mature ova without pigmented poles; nuptial pads, vocal sac, vocal slits, enlarged forearms, all absent. Morphological comparison. Characters used for comparing Megophrys periosa sp. nov. with its congeners do not include those from the provisionally assigned referred specimens. Megophrys periosa sp. nov. (adult males, N =12, adult female, N =1) differs from M. medogensis, M. zhangi and M. monticola by its larger adult body size, male SVL 71.3–93.8 mm, female SVL 112.0 mm (vs. male SVL 57.2–68.0 mm, N =17; male SVL 32.5–37.2 mm, N =3; male SVL 38.2–49.5 mm, N =17, female SVL 40.5–56.1 mm, N =6, respectively); differs from M. robusta by absence of black dermal asperities on posterior abdomen of adult males (vs. usually present), Finger IIN =13 (vs. Finger I=II, N =10); differs from Megophrys flavipunctata sp. nov., Megophrys oreocrypta sp. nov., M. major s.s. and M. mangshanensis by absence of distinct white, cream or light coloured stripe along upper lips (vs. present); further differs from Megophrys flavipunctata sp. nov. and M. mangshanensis by its larger adult body size, male SVL 71.3–93.8 mm, female SVL 112.0 mm (vs. male SVL 56.9–68.4 mm, N =4, female SVL 68.0– 74.6 mm, N =3; male SVL 62.5 mm, N =1, female SVL 73 mm, N =1, respectively); further from M. major s.s. by dark spots associated with flank tubercles typically present (vs. absent), toe tips not expanded relative to adjacent toe width (vs. expanded); differs from Megophrys himalayana sp. nov. by its typically larger adult body size, male SVL 71.3–93.8 mm, female SVL 112.0 mm (vs. male SVL 68.0– 73.5 mm, N =7, female SVL 83.9 mm, N =1), typically smaller eye diameter to snout length ratio for males, ED/ES 76.6–91.5%, mean 84.3 ± 4.2% (vs. ED/ES 87.6–101.1%, N =7, mean 94.2 ± 4.6%). Systematic position. This taxon represents M. cf. major 4 (OTU 12) in the molecular analyses, and “ M. cf. major [5]” in Mahony et al. (2017). Megophrys periosa sp. nov. consistently formed a sister taxa relationship with two populations found east of the Brahmaputra River in Manipur and Myanmar referred to herein as Megophrys cf. periosa (M. cf. major 6/OTU 14 in molecular analyses). Uncorrected p -distances for the 16S rRNA gene between Megophrys periosa sp. nov. and M. cf. periosa was 1.6–2.0% (Appendix I, Table 6). The systematic position of this clade within the MMC was not fully resolved (see Systematic position section for Megophrys himalayana sp. nov. for further details), however most analyses placed this species as the sister taxon of Megophrys himalayana sp. nov. (Figures 2 & 4; Appendix I, Table 3; Appendix II, Figures 1, 2 & 5; Mahony et al. 2017). Refer to Appendix I, Table 6 for uncorrected p - distances for the 16S rRNA gene between Megophrys periosa sp. nov. and other MMSG species. Etymology. The species epithet “ periosa ” is a Latinised Greek adjective meaning “immense”, in reference to the very large size that this species attains, even compared to otherwise generally large species in the MMC. Suggested common name: Giant Himalayan Horned Frog. Distribution. Megophrys periosa sp. nov. is so far confirmed from three localities in the state of Arunachal Pradesh, Northeast India. The currently known distribution ranges from East Siang district, west through West Siang district, to at least West Kameng district, between 260 and 1110 m asl. (Figure 8A). An improved and more extensive sampling is necessary to define the east-west limits of this species’ geographic range. Specimens assigned here to Megophrys cf. periosa were collected from Ukhrul district (at 820 m asl.), Manipur state, Northeast India, and from Myitkyina district, Kachin state, northern Myanmar (Figure 8A). Habitat and natural history. All males were in breeding condition, and the female contained well-developed ova within her ovaries, indicative that the breeding season extends at least through late-July and early-August. All specimens were collected after dusk, typically perched on the rocky banks of small to large (1–15 m wide) moderately fast flowing mountain streams flowing through dense mature secondary/primary forest (e.g., Figure 9A & F). Males spaced themselves along the banks of the streams at least 10 m apart. Vocalisations typically consisted of a succession of a few notes followed by an extended silence, but were rarely heard. Healed injuries were observed on two specimens; the right hand of the holotype appears to have been badly dislocated or broken (Figure 22B & E). This large mature male had noticeably less well-developed nuptial pads than the other males, possibly as a result of lower overall fitness, although it otherwise appeared to be in good health. One other specimen (BNHS 6064) had a large swelling on Finger II on the right hand, the cause for which is unclear. Remarks. Specimens reported as M. lateralis and/or M. robusta by Borah and Bordoloi (2001), Bordoloi et al. (2000) and Sarkar and Ray (2006) from Arunachal Pradesh may apply to this species, and/or Megophrys himalayana sp. nov. The specimens reported in these studies should be re-examined to determine the correct identities. Populations provisionally assigned to this taxa (as M. cf. periosa) from eastern Manipur state, Northeast India and northern Myanmar, were found to be the sister taxon to Megophrys periosa sp. nov. (Mahony et al. 2017 [as M. cf. major [6]]; Figures 2, 4 & 5). Regardless of BPP results indicating that this lineage may represent a distinct species-level taxon (Appendix I, Table 5), we recommend that comparable series of adult specimens be studied to further determine the taxonomic status of this lineage. For now, it may provisionally be regarded as conspecific with Megophrys periosa sp. nov.

Megophrys (Xenophrys) flavipunctata sp. nov. (Figures 16 & 17; Table 1) Holotype. Adult male (BNHS 6040 [field no. SDBDU 2009.304]: Figures 16 & 17B), from Mawphlang Sacred Grove, Mawphlang (25��26'37"N, 91��44'46"E, 1810 m asl.), East Khasi Hills district, Meghalaya state, Northeast India, collected by members of the Systematics Lab, University of Delhi on 0 1 June 2009. Paratypes. One adult male (BNHS 6044 [field no. SDBDU 2009.305]), and two adult females (BNHS 6043 [field no. SDBDU 2009.303]: Figure 17A, C & D; BNHS 6042 [field no. SDBDU 2009.306]: Figure 18A), collected along with the holotype; one adult female (BNHS 6041 [field no. SDBDU 2011.480]: Figure 17A), collected from the type locality by SDB and RGK on 0 9 July 2011. Referred specimens. One adult male (SDBDU 2009.297), and one juvenile male (SDBDU 2009.298), collected from the type locality by Systematics Lab members on 30 May 2009; one adult male (SDBDU 2007.134: Figure 17E & F), from near Sazolie College, Jotsoma village (25��38'23"N, 94��03'31"E, 1575 m asl.), Kohima district, Nagaland state, Northeast India, collected by RGK on 27 June 2007. Holotype description (measurements in mm). Mature male (SVL 68.4) (Figures 16 & 17B). Head moderately large, wider than long (HW 26.3, HL 25.0, IFE 10.5, IBE 18.1); snout bluntly pointed in dorsal view, obtusely protruding beyond mandible in lateral view, without rostral appendage (Figure 16C); loreal region acute, concave; canthus rostralis angular; dorsal surface of snout slightly concave; eye diameter 2.5 times maximum diameter of visible portion of tympanum, and shorter than snout (EL 7.6, TYD 3.1, SL 8.6); eye���tympanum distance (TYE 5.8) longer than diameter of visible portion of tympanum; tympanum oval-shaped, slightly oblique, upper border concealed by supratympanic ridge (Figure 16C); pupil in life vertically elliptical; nostril positioned laterally, closer to eye than to snout (EN 3.8, NS 5.4); internarial distance greater than eyelid width and narrowest point between upper eyelids (IN 7.7, UEW 6.8, IUE 5.4); pineal ocellus not visible externally; vomerine ridges medium sized, ovoid, moderately raised, orientated acutely, positioned between to slightly posterior to choanae, equidistant from each other and choanae; vomerine teeth small; maxillary teeth present; tongue moderately large, deeply notched posteriorly, medial lingual process absent. Forelimbs moderately long, thin (Figure 16A & B), forearm not enlarged relative to upper forelimb, and shorter than hand (FAL 14.6; HAL 17.9); fingers moderately long, narrow, without lateral fringes (Figure 16D), finger length formula IV Colouration: In preservative (Figure 16): Dorsal and lateral surfaces of head and body primarily mid greyishbrown mottled with lighter tones; light-edged, dark brown triangular marking present between eyes; short longitudinal brown stripe on mid-dorsal snout; small light-edged dark brown Y-shaped marking on anterior dorsum, dark brown, ragged-edged, dorsolateral stripe present on each side; tubercles on anterior flanks dark brown with light tip, on posterior flanks pale yellow with bordering small dark brown spot; front of snout greyishbrown with extensive yellowish-orange speckling, lateral canthus rostralis dark greyish-brown; light grey stripe on upper lip extends from nostril to rear of upper jaw, bordered below by thin dark brown stripe along margin of lip; dark brown blotch extends from posterior border of eye through tympanum to posterior supratympanic ridge; additional triangular blotch distinct at posterior lower orbital border, with intervening area a lighter shade of brown; outer edges of eyelids dark brown; lower half of supratympanic ridges dark brown, upper half light yellowish-cream which continues anteriorly as fine light stripe along edge of upper eyelids and canthus rostralis to tip of snout; dorsal surfaces of forelimbs and hindlimbs primarily light greyish-brown with mid brown blotches; two dark brown blotches on anterior lateral surface of forearms; dorsal surface of outer three fingers with dark brown blotches; dorsal surfaces of hindlimbs with distinct mid brown transverse crossbars; lateral surfaces of thighs and shanks with dark brown spots and blotches; throat primarily light brown with some yellow spots along its edge, light-edged wide dark brown stripe extends from posterior edge of lower jaw to ventral forearm; chest and anterior abdomen with faint large mid brown blotches on otherwise pale yellowish-grey background; ventral surfaces of forelimbs and shanks primarily pale grey, thighs pale yellowish-grey scattered with minute yellow spots; some large dark brown spots and blotches laterally along abdomen on both sides; area surrounding cloaca and posterior surfaces of thighs primarily dark brown with small yellow spots; ventral surfaces of tarsi and feet greyish-brown; hands ventrally light grey; pectoral and femoral glands creamish-white. In life (Figure 17B): Colouration mostly as described in preservative but richer; enlarged tubercles on posterior and ventrolateral flanks, and enlarged portion of posterior supratympanic ridge bright yellow; iris colour rich maroon. Ventral colouration in life not documented. Variation. See Table 1 for morphometric variation within the type series and referred specimens consisting of three adult males, three adult females, and a juvenile. The paratypes and referred specimens resemble the holotype for most morphological characters with some exceptions: BNHS 6041, BNHS 6043 and SDBDU 2009.298 do not have distinct lateral fringes on toes; BNHS 6044 has finger length formula IISecondary sexual characters. Males: weakly raised nuptial pads present, sparsely covered with black microasperities, primarily covering dorsal surface of base of Finger I, narrowing distally, extending onto base of distal phalange on inner dorsal side; nuptial pad on Finger II small, oval, positioned on base of digit on inner dorsal side; external vocal sac indistinct; internal vocal slits present, one on each side on floor of mouth near rear of mandible; forearms not enlarged relative to upper forelimbs. Females: mature ova not pigmented; nuptial pads, vocal sac, vocal slits, enlarged forearms, all absent. Morphological comparison. Megophrys flavipunctata sp. nov. (adult males, N =4, adult females, N =3) differs from M. monticola and M. zhangi by its larger adult body size, male SVL 56.9���68.4 mm, female SVL 68.0��� 74.6 mm (vs. male SVL 38.2���49.5 mm, N =17, female SVL 40.5���56.1 mm, N =6; male SVL 32.5���37.2 mm, N =3, respectively); differs from M. medogensis by presence of vivid whitish-cream upper lip stripe (vs. absent), toes dorsoventrally flattened, with or without narrow lateral dermal fringes (vs. toes rounded, not flattened, lateral fringes absent); differs from M. robusta and M. major s.s. by smaller adult body size, male SVL 56.9���68.4 mm, female SVL 68.0��� 74.6 mm (vs. male SVL 73.5���83.1 mm, N =6, female SVL 81.3���108.3 mm, N =4; male SVL 71.6���87.5 mm, N =12, female SVL 85.6���98.2 mm, N =3, respectively), further from M. robusta by presence of vivid whitish-cream upper lip stripe (vs. absent), further from M. major s.s. by distinct brown blotches clearly visible on chest and anterior abdomen of all specimens (vs. typically absent, if present always very faint), flank tubercles, particularly the anteriormost tubercles with dark brown blotches present (vs. absent); differs from M. mangshanensis by basal webbing between toes present (vs. absent). For comparisons with subsequent species covered in this study, refer to relevant morphological comparison sections for those species. Systematic position. This taxon represents M. cf. major 2 in the molecular analyses of this study (discussed as OTU 6), and in Mahony et al. (2017). The systematic position of Megophrys flavipunctata sp. nov. within the MMC clade is not fully resolved. It has consistently been found to be a member of the western clade of the MMC (Figure 1), however, its relationship with other members of this clade is not clear when molecular data for the MMSG was subjected to a variety of phylogenetic techniques (Figures 2���5; Appendix I, Table 3; Appendix II, Figures 1, 2, 5 & 6). Refer to Appendix I, Table 6 for uncorrected p -distances for the 16S rRNA gene between Megophrys flavipunctata sp. nov. and other MMSG species. Etymology. The specific epithet is an adjective derived from the Latin words flavus meaning ���yellow��� and punctatus meaning ���spotted��� with reference to the bright yellow tubercles on the posterior flanks of this species. Suggested common name: Yellow Spotted White-lipped Horned Frog. Distribution. Megophrys flavipunctata sp. nov. is currently known with certainty from two populations, one in Meghalaya state, and a second disjunct population in Nagaland state, Northeast India (Figure 8A). Mawphlang Sacred Grove (25��26'37.02"N, 91��44'46.02"E, 1810 m asl.) is located next to the village of Mawphlang, on the upper limits of the Shillong Plateau, southwest from Shillong city, Meghalaya. The second known population is represented by a single specimen collected from near Sazolie College, Jotsoma village (25��38'23"N, 94��03'31"E, 1575 m asl.), Kohima district, Nagaland state, ~ 230 km east of the type locality. This species is likely to be present at other mid- to high-elevation localities in intervening areas, where suitable habitat remains. It may also extend westward in Meghalaya to Tura Peak, West Garo Hills district (see Remarks below for Sangma and Saikia 2015), however this locality requires further confirmation. Habitat and natural history. Refer to Mahony (2008) and Mahony et al. (2013) for a general description of the habitat and forest type at Mawphlang (Figure 9B & D). The type series was collected on the forest floor, on or amongst dense leaf litter. Those collected at dusk were found emerging from beneath leaf litter, and one from the entrance of a rodent burrow, where presumably this species remains concealed during daylight hours. Two other individuals collected immediately after dusk, were on top of leaf litter several meters to the side of a forest trail in a part of the forest with sparse to no undergrowth. All were more than 50 meters from the nearest stream. Despite numerous visits to Mawphlang Sacred Grove between May and September 2006 to 2011, this species was only observed in 2009 and 2011, possibly because earlier surveys were primarily focused along streams and forest trails. No male advertisement calls attributable to this species were heard on any visit. This species was also not reported by others carrying out amphibian research at Mawphlang (e.g., Das et al. 2010; Purkayashta & Matsui 2010; Tron et al. 2015). The distant location of adult animals from streams in late May/early June might indicate that this species does not breed at this time of the year. Adult females had moderately well developed ova in their ovaries, and males also had moderately small (but developmentally mature) testes and very weakly developed nuptial pads indicating that they were not in full reproductive condition. Further field studies are required to determine the breeding season. The single specimen from Nagaland was collected in the month of June, on a road (at 22:30 h), adjacent to a temporary stream, nearby a degraded semi-deciduous forest patch (see Biju et al. 2010 for discussion on habitat condition at this locality). This adult male was also not in breeding condition, i.e., nuptial pads not developed, and testes not enlarged. Remarks. A previous report of M. glandulosa from Puliebadze (near Jotsoma), Nagaland by Ao et al. (2003) might represent this species (see Remarks section of M. glandulosa). Sangma and Saikia (2015:2410, pl. 6a) provided a figure of a specimen that superficially resembles this new species (as Xenophrys robusta) from Tura Peak (751 m asl.), West Garo Hills, Meghalaya, however the identity of this specimen requires further confirmation pending examination., Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on pages 46-50, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/2610202, {"references":["Mahony, S., Foley, N. C., Biju, S. D. & Teeling, E. C. (2017) Evolutionary history of the Asian Horned Frogs (Megophryinae): integrative approaches to timetree dating in the absence of a fossil record. Molecular Biology and Evolution, 34 (3), 744 - 771 + S. I. 33 pp. https: // doi. org / 10.1093 / molbev / msw 267","Sangma, M. A. & Saikia, P. K. (2015) Checklist of amphibians inventoried from Tura Peak of West Garo Hills, Meghalaya, India. Journal of Global Biosciences, 4 (5), 2404 - 2424.","Mahony, S. (2008) Redescription and generic reallocation of Rana mawphlangensis Pillai & Chanda, 1977 (Amphibia: Ranidae). Hamadryad, 33 (1), 1 - 12.","Mahony, S., Teeling, E. C. & Biju, S. D. (2013) Three new species of horned frogs, Megophrys (Amphibia: Megophryidae), from Northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region. Zootaxa, 3722 (2), 143 - 169. https: // doi. org / 10.11646 / zootaxa. 3722.2.2","Das, I., Tron, R. K. L., Rangad, D. & Hooroo, R. N. (2010) A new species of Leptolalax (Anura: Megophryidae) from the sacred groves of Mawphlang, Meghalaya, north-eastern India. Zootaxa, 2339, 44 - 56.","Purakayastha J. & Matsui M. (2010) A new species of Fejervarya (Anura: Dicroglossidae) from Mawphlang, northeastern India. Asian Herpetological Research, 3 (1), 31 - 37.","Tron, R. K. L., Das, I., Hooroo, R. N. K. & Rangad, D. (2015) Spring-breeding and reproductive mode in Leptolalax khasiorum (Anura, Megophryidae) in northeastern India. Russian Journal of Herpetology, 22 (3), 224 - 232.","Biju, S. D., Mahony, S. & Kamei, R. G. (2010) Description of two new species of torrent frog, Amolops Cope (Anura: Ranidae) from a degrading forest in the northeast Indian state of Nagaland. Zootaxa, 2408, 31 - 46.","Ao, M., Bordoloi, S. & Ohler, A. (2003) Amphibian fauna of Nagaland with nineteen new records from the State including five new records for India. Zoos' Print Journal, 18 (6), 1117 - 1125. https: // doi. org / 10.11609 / JoTT. ZPJ. 18.6.1117 - 25"]}

Megophrys (Xenophrys) medogensis Fei, Ye and Huang, 1983 (Figure 13) Megophrys omeimontis medogensis Fei, Ye and Huang 1983:49 In: Two new subspecies of Megophrys omeimontis Liu from China (Amphibia, Pelobatidae). Acta Herpetologica Sinica, New Series, 2(2): 49���52. Holotype. Adult male (CIB 73 II0015: Figure 13), from ���Motuo, Xizang, alt. 1000m ��� (= Medog [or Motuo] town [~ 29��19'56"N, 95��20'27"E], Medog County, Nyingtri Prefecture, Tibet [or Xizang] Autonomous Region, China), collectors presumably Fei, Hu, Ye and Wu on 17 July 1973 (Fei et al. 1977). Paratypes [not examined]. 16 adult males, one juvenile and some tadpoles (CIB specimen numbers not provided), ���collected from the same locality, alt. 850���1350m ���, collectors as above, including Huang, on 17���24 July 1973 and 21 July���5 August 1977 (Fei et al. 1977). Examined specimens. Adult male (CIB 73II 0015: images only: Figure 13), holotype. Holotype description. Refer to Fei et al. 1983 for the holotype description. For additional details of colouration and markings, see figures in Zhao et al. (2005), Fei et al. (2010, 2012) and Li et al. (2010) of specimens from the type locality. Systematic position. Refer to the Systematic position section above for M. robusta. Morphological comparison. Adult body size ranges provided for M. medogensis are from Fei et al. (2009) (adult males, N =16). Megophrys medogensis differs from M. monticola and M. zhangi by its much larger adult body size, male SVL 57.2���68.0 mm (vs. male SVL 38.2���49.5 mm, N =17; male SVL 32.5���37.2 mm, N =3, respectively); differs from M. mangshanensis by absence of white upper lip stripe (vs. present); differs from M. robusta by its smaller adult male size, SVL 57.2���68.0 mm (vs. male SVL 73.5���83.1 mm, N =6). For comparisons with additional species covered in this study, refer to relevant morphological comparison sections for those species. Etymology. The specific epithet ��� medogensis ��� is a toponym, derived from the type locality of Medog county. Suggested common name: Medog Horned Frog. Distribution. Megophrys medogensis is currently known with certainty from the type locality (Figure 8A) ���Medog���, between 850 and 1350 m elevation (Fei et al. 1983, 2012). This species has also been reported from the southern slopes of Mt. Namjagbarwa (or Namcha Barwa) adjacent to Medog town (Zhao & Li 1987). Additional localities in Yunnan Province, China on the distribution map provided by Fei et al. (2009) are erroneous (see Remarks below). The type locality is situated near to the northern border of the Upper Siang district, Arunachal Pradesh state, Northeast India, where this species is likely to be present at similar elevation. Habitat and natural history. At the type locality, males were reported to frequent leaf litter and small streams in forests, and were heard vocalising especially on nights with clear skies (14 July���08 August). Tadpoles allocated to this species were collected from amongst rocks at the base of a waterfall of a small stream (Fei et al. 2009), but the authors did not justify how they identified the tadpoles. Remarks. In the English abstract of the original description, Fei et al. (1983) only provided a brief comparison of this species with M. omeimontis Liu, 1950 and M. jingdongensis Fei and Ye, 1983 (in Fei et al. 1983), congeners that they determined to be morphologically most similar to M. medogensis. The abstract provided the name as ��� Megophrys omeimontis motuoensis ���, but elsewhere in the text the name was provided as ��� Megophrys omeimontis medogensis ���, indicating that the name in the abstract was an accidental erroneous spelling (also noted by Zhao & Adler 1993). Fei et al. (1983) provided a line drawing of the profile view of the head and ventral view of the foot, but not measurements of specimens. Fei et al. (2009) provided a table of measurements for the type series, and Fei et al. (2010) provided low-resolution photos of the holotype in preservation. Zhao et al. (2005), Fei et al. (2010, 2012), and Li et al. (2010) provided images of live individuals. Fei et al. (2009) included a distribution map showing several localities in Yunnan province for Megophrys medogensis but did not provide their source (neither publications nor specimens) for these additional localities. In Fei et al. ���s (2009) book, the species Megophrys jingdongensis is accompanied by a distribution map that is identical to the one provided for M. medogensis, but without the locality ���Medog��� marked. Thus it is likely that the Yunnan localities for M. medogensis are the result of printing error, and should not be considered for this species��� distribution. Fei et al. (2012) provided a map for this species including only the type locality, indirectly correcting the error of Fei et al. (2009)., Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on pages 37-39, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/2610202, {"references":["Fei, L., Ye, C. - Y. & Huang, Y. - Z. (1983) Two new subspecies of Megophrys omeimontis Liu from China (Amphibia; Pelobatidae). Acta Herpetologica Sinica, 2 (2), 49 - 52.","Fei, L., Ye, C. Y., Wu, G. F. & Hu, S. Q. (1977) A survey of amphibians in Xizang (Tibet). Acta Zoologica Sinica, 23 (1), 54 - 63.","Zhao, W, Rao, D., Lu, S. & Dong, B. (2005) Herpetological surveys of Xizang Autonomous Region. 2. Medog. Sichuan Journal of Zoology, 24 (3), 250 - 253.","Fei, L., Ye, C. & Jiang, J. (2010) Coloured Atlas of Chinese Amphibians. Sichuan Publishing House of Science & Technology, Sichuan, 517 pp.","Fei, L., Ye, C. - Y. & Jiang, J. - P. (2012) Coloured Atlas of Chinese Amphibians and their Distributions. Sichuan Publishing House of Science & Technology, Sichuan, 620 pp.","Li, P. - P., Zhao, E. - M. & Dong, B. - J. (2010) Amphibians and Reptiles of Tibet. Science Press, Beijing, XII + 251 pp.","Fei, L., Hu, S. - Q., Ye, C. - Y. & Huang, Y. - Z. (2009) Fauna Sinica. Amphibia. Folume 2. Anura. Chinese Academy of Science, Science, Beijing, XIII + 957 pp.","Zhao, E. & Li, S. (1987) The herpetofaunal composition and characteristics of Mount Namjagbarwa, Tibet. Acta Herpetologica Sinica, 6 (2), 36 - 42.","Liu, C. - C. (1950) Amphibians of western China. Fieldiana, 2, 1 - 400, pls. 1 - 10.","Zhao, E. - m. & Adler, K. (1993) Herpetology of China. SSAR, Oxford, 552 + V pp., pls. 1 - 48 + 1."]}

Megophrys (Xenophrys) glandulosa Fei, Ye and Huang, 1990 (Figure 24; Table 1) Megophrys glandulosa Fei, Ye and Huang 1990:99, 273. In: Key to Chinese Amphibians. Publishing House for Scientific and Technological Literature, Chongqing, China: 364 pp. + 2 + [iv]. Holotype. Adult male (CIB 873112: Figure 24), from “Wuliang shan, Jingdong, Yunnan Province, altitude 1900m ” (= Wuliang Mountain [~ 24°49'27"N, 100°26'31"E], Jingdong Yi Autonomous County, Pu’er Prefecture, Yunnan Province, China), collected by Liang Fei on 29 March 1987 (Fei et al. 1992). Paratypes. Adult female (“ allotype ” CIB 873201), from the “same locality as that of the holotype, altitude 2100m ”, collector not mentioned, 0 2 May 1975; 25 males and two females (specimen numbers not mentioned in Fei et al. [1992]), from “Wuliang shan, Jingdong, Yunnan Province, altitude 1900–2100m ”, collectors Liang Fei, Younzhao Huang and Jiarui Luo, between 29 and 30 March 1987 (Fei et al. 1992). Examined specimens. Adult male (CIB 873112––photos only), holotype. Provisionally referred examined specimens. Two adult males (CAS 221395, CAS 221484), from two localities on the road between Ahtonga and Babaw (27°15'27.2"N, 97°50'32.4"E and 27°17'24.3"N, 97°51'52.4"E), Machanbaw township, Putao district, Kachin state, Myanmar, collected by H. Win and R. Shaung on 0 5 September 2001, and J.B. Slowinski, D.-O. Rao, G.O.U. Wogan, H. Win, A.K. Shein and H. Tun on 0 7 September 2001; one adult male (CAS 221442), from Aureinga Camp (27°17'36.4"N, 97°51'50"E), Naung Mon township, Putao district, Kachin state, Myanmar, collected by J.B. Slowinski, G.O.U. Wogan, H. Win and A.K. Shein on 0 7 September 2001. Description of type series. Refer to Fei et al. (1992) for the description in Chinese language text, and Huang et al. (1998) for the English translation of Fei et al. (1992). See Figure 24 for colouration in preservative of the holotype. Description of referred specimen CAS 22 1442 (measurements in mm). Mature male (SVL 81.0). Head large (HW 31.3, HL 30.2, IFE 13.6, IBE 21.2), wider than long, snout broadly pointed in dorsal view, obtusely protruding beyond mandible in lateral view, without rostral appendage; loreal region acute, concave, with well developed canthus rostralis; dorsal surface of snout concave; eye diameter larger than maximum tympanum diameter, and shorter than snout (EL 7.9, TYD 5.1, SL 10.1); eye–tympanum distance (TYE 6.7) longer than tympanum diameter; tympanum distinctly oval, obliquely orientated, upper ~10% concealed by supratympanic ridge; pupil vertically elliptical; nostril positioned laterally, oval, obliquely oriented with raised posterior rim, closer to eye than to snout (EN 4.6, SN 6.2); upper eyelid width subequal to narrowest point between upper eyelids, and less than internarial distance (UEW 7.5, IUE 7.6, IN 9.3); pineal ocellus absent; vomerine ridges well developed, ovoid, positioned between choanae, extending posterior to choanae, slightly closer to choanae than to each other; vomerine teeth present, moderately long; maxillary teeth present; tongue moderately large, distinctly notched posteriorly, medial lingual process absent. Forelimbs moderately long, thick; forearms enlarged relative to upper forelimbs, and shorter than hand length (FAL 17.3, HAL 21.3); fingers moderately long, narrow, finger length formula IVColouration: In preservative: Dorsal and lateral surfaces of head and body primarily plain brown; light-edged, dark brown triangular marking between eyes; faint darker brown V-shaped marking on anterior dorsum surrounding parietoscapular ridge, narrow broken dark brown stripe follows lower edge of dorsolateral ridges; larger tubercles on flanks dark brown with light tip; front of snout and lateral surfaces below canthus rostralis dark brown; greyish-white stripe on upper lip extends from nostril to rear of upper jaw, interrupted by wide dark brown vertical bar between lower edge of orbits and edge of mouth; besides narrow brown oblique stripe extending from posterior edge of orbits posteriorly to join light upper lip stripe, remaining lateral surfaces of head below supratympanic ridges solid dark brown; outer edges of eyelids dark brown; lower half of supratympanic ridges dark brown, upper half brown anteriorly, light yellowish-cream posteriorly; dorsal surfaces of forelimbs and hindlimbs brown; two dark brown blotches on anterior lateral surface of forearms; dorsal surface of outer three fingers with dark brown blotches; dorsal surfaces of hindlimbs with distinct brown transverse crossbars; outer lateral surfaces of thighs and shanks with dark brown spots and blotches; throat with large light brown blotches, few small white spots along its edge, light-edged wide dark brown stripe extends from posterior edge of lower jaw to ventral surface of upper forelimb on each side; chest and abdomen with faint large brown blotches on an otherwise pale greyishwhite background; ventral surfaces of forelimbs pale greyish-white; ventral surfaces of thighs densely mottled light brown and greyish-white, posterior surfaces of thighs (including area surrounding cloaca) and ventral surfaces of shanks and tarsi primarily mid to dark brown with pale greyish-white scattered spots; some large dark brown spots and blotches laterally along abdomen on both sides; ventral surfaces of hands and feet greyish-brown; pectoral and femoral glands creamish-white. In life: Not documented for this specimen. Variation. Refer to Table 1 for morphometric variation within the provisionally referred specimen series, consisting of three adult males. The two other referred specimens generally resemble the referred specimen described above for most morphological characters with the following exceptions: CAS 221484 has more extensive webbing between Toes II to V (i.e., webbing formula: II2 –3.1 III2.8 – 4 IV 4– 2.5 V); CAS 221484 has relative finger lengths II=ISecondary sexual characters. Males: nuptial pads present, weakly raised, covered with black micro-granules, covering almost entire dorsal surface of Finger I narrowing distally where it extends to base of distal phalange on inner dorsal side; nuptial pad small to moderately large on Finger II, positioned on base of digit on inner dorsal side (extending on to base of distal phalange on some individuals); external vocal sac indistinct; internal vocal slit present on floor of mouth on each side near rear of mandible; forearms enlarged relative to upper forelimbs. Females: not examined in this study. Morphological comparison. Adult body size ranges provided for Megophrys glandulosa include the measurements given in Fei et al. (2009) and the referred specimens examined in this study (total adult males, N =13; adult females, N =3). The remaining characters were assessed based only on the referred specimens and images of the holotype (total: adult males, N =4). Megophrys glandulosa differs from M. medogensis, M. zhangi, M. monticola, Megophrys periosa sp. nov. and M. robusta by the presence of a distinct white upper lip stripe (vs. absent), presence of broad lateral fringes on toes (vs. absent on M. medogensis, M. zhangi, M. monticola, and M. robusta; usually absent, though very narrow when present on Megophrys periosa sp. nov.), and differs further from M. medogensis, M. zhangi, and M. monticola by its larger adult size, male SVL 76.3–81.0 mm, female SVL 76.5–99.5 mm (vs. male SVL 57.2–68.0 mm, N =16; male SVL 32.5–37.2 mm, N =3; male SVL 38.2–49.5 mm, N =17, female SVL 40.5–56.1 mm, N =6, respectively); differs from Megophrys himalayana sp. nov. and M. mangshanensis by its larger adult male size, SVL 76.3–81.0 mm (vs. male SVL 68.0– 73.5 mm, N =7; male SVL 62.5 mm, N =1, respectively), and by broad lateral fringes on toes present (vs. absent); differs from M. major s.s. and Megophrys oreocrypta sp. nov. by presence of broad lateral fringes on toes, and toe tips not expanded (vs. occasional presence of narrow lateral fringes on toes, and toe tips distinctly expanded relative to toe width), ventral surface of thighs densely spotted and speckled (vs. immaculate); differs from Megophrys flavipunctata sp. nov. by its larger adult size, male SVL 76.3–81.0 mm, female SVL 76.5– 99.5 mm (vs. male SVL 56.9–68.4 mm, N =4, female SVL 68.0– 74.6 mm, N =3), by toe tips not expanded relative to toe width (vs. toe tips slightly expanded), ventral thighs densely spotted and speckled (vs. immaculate). Systematic position. The systematic position of Megophrys glandulosa within the MMC could not be resolved in this study possibly due to incomplete sampling of nuclear markers (i.e., absence of CXCR4, TTN and SACS; Appendix I, Table 2). Analyses of the mtDNA datasets (16S and CO1 gene trees, and 16S+CO1 concatenated tree), the RHOD gene, and the concatenated mt+nuDNA dataset indicated that this species is a distinct lineage (Figure 4; Appendix II, Figure 5), however, the RAG1 gene tree (not shown) and concatenated nuDNA analyses indicated that this species may have close affinity to the species referred to here as M. cf. maosonensis [1], though this relationship received very low support (Appendix II, Figure 6). Etymology. The specific epithet “ glandulosa ” is Latin, with glandul meaning “gland”, and the suffix - osa meaning “abundant”, presumably a reference to the tubercles on the flanks of this species, a character that is not useful for diagnosing this species from other species in the MMC. Suggested common name: Glandular White-lipped Horned Frog. Distribution. Besides the type locality, M. glandulosa has been reported from numerous localities in Yunnan Province between 1900 and 2500 m asl. (e.g., Jiang et al. 2003; Fu et al. 2006; Fei et al. 2009), and Kachin state, northern Myanmar by Wogan et al. (2008) (Figure 8A). Reports of this species from further west in Northeast India and Bhutan are not here considered reliable (see Remarks below). Remarks. Megophrys glandulosa was described (in Chinese text) as new twice based on the same type series (Fei et al. 1990, 1992). Fei et al. (1992) was subsequently translated into English by Huang et al. (1998). It is highly unlikely that the Bhutan (Wangyal 2013) and Indian (Meghalaya, Nagaland and West Bengal states: Ao et al. 2003; Deuti & Ayyaswamy 2008; Mathew & Sen 2010; Sangma & Saikia 2015) populations are conspecific with Megophrys glandulosa s.s. Ao et al. (2003) reported a specimen of this species from Puliebadze (25°37'0"N, 94°5'0"E, 2225 m asl.), Kohima district, Nagaland state, Northeast India. Though we have not examined this specimen, we report two species of MMC from Kohima district in this study (M. major s.s. 450– 1255 m asl. and Megophrys flavipunctata sp. nov. 1575–1810 m asl.). Based on the elevation that Ao et al.’s (2003) specimen was collected, we suspect that it might represent Megophrys flavipunctata sp. nov. Mathew and Sen (2010) provided an undiagnostic description of M. glandulosa for India with an accompanying photo of a specimen that appears to represent M. major s.s. A report of two specimens of Megophrys [as Xenophrys] glandulosa from Tura Peak, West Garo Hills, in Meghalaya state provided by Sangma and Saikia (2015: 2407, pl. 4a) appears to represent an adult Megophrys oreocrypta sp. nov. based on the figure provided for one of the specimens. An additional report of M. glandulosa by Deuti and Ayyaswamy (2008) was based on a collection from Latpanchar, Darjeeling district, West Bengal state, Northeast India. The figured individual displays typical characters seen on M. robusta, i.e., slightly enlarged posterior supratympanic ridge (vs. greatly enlarged, and beanshaped on M. glandulosa [Figure 24C]); the distinctive white upper lip stripe of M. glandulosa is not apparent on the figured animal, and the iris colour matches that of M. robusta perfectly. Although some of the characters that were described, such as ventral markings, could correspond with M. glandulosa, other characters such as swollen digit tips, and rudimentary webbing between toes do not (see description provided above). Pending re-examination of the specimens reported by Deuti and Ayyaswamy (2008), we recommend that Darjeeling, and Northeast India in general, are not included in the distribution of M. glandulosa. Wangyal (2013) reported M. glandulosa and M. major from Bhutan, however, the two specimens photographed exhibit characters that allow a reasonable preliminary identification as M. robusta, i.e., both lack the light upper lip stripe seen on M. glandulosa and M. major, and have pale iris colouration that is characteristic of the M. robusta / M. medogensis clade in the MMSG., Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on pages 64-67, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/2610202, {"references":["Fei, L., Ye, C. - Y. & Huang, Y. - Z. (1990) Key to Chinese Amphibians. Chongqing Branch, Science and Technology Literature Publishing House, Chongqing, 364 + 2 + [iv] pp.","Fei, L., Ye, C. - Y. & Huang, Y. - Z. (1992) Two new species of the genus Megophrys, Pelobatidae (Amphibia, Anura) from China. Zoological Research, 13 (1), 5 - 12.","Huang, C. - M., Lathrop, A. & Murphy, R. W. (1998) Translations of \" Two new species of amphibian in Tibet \" Huang & Fei 1981 and \" Description of two new species of the genus Megophrys \" Fei, Ye, & Huang 1995 [1992]. Smithsonian Herpetological Information Service, 118, 1 - 20.","Fei, L., Hu, S. - Q., Ye, C. - Y. & Huang, Y. - Z. (2009) Fauna Sinica. Amphibia. Folume 2. Anura. Chinese Academy of Science, Science, Beijing, XIII + 957 pp.","Jiang, J. P., Yuan, F. R., Xie, F. & Zheng, Z. H. (2003) Phylogenetic relationships of some species and genera in megophryids inferred from partial sequences of mitochondrial 12 S and 16 S rRNA genes. Zoological Research, 24 (4), 241 - 248.","Fu, C., Hua, X., Li, J., Chang, Z., Pu, Z. & Chen, J. (2006) Elevational patterns of frog species richness and endemic richness in the Hengduan Mountains, China: geometric constraints, area and climate effects. Ecography, 29, 919 - 927 + 1 - 7 (Appendix E 4802).","Wogan, G. O. U., Vindum, J. V., Wilkinson, J. A., Koo, M. S., Slowinski, J. B., Win, H. Thin, T. Kyi, S. W., Oo, S. L., Lwin, K. S. & Shein, A. K. (2008) New country records and range extensions for Myanmar amphibians and reptiles. Hamadryad, 33, 83 - 96.","Wangyal, J. T. (2013) New records of reptiles and amphibians from Bhutan. Journal of Threatened Taxa, 5, 4774 - 4783. https: // doi. org / 10.11609 / JoTT. o 3539.4774 - 83","Ao, M., Bordoloi, S. & Ohler, A. (2003) Amphibian fauna of Nagaland with nineteen new records from the State including five new records for India. Zoos' Print Journal, 18 (6), 1117 - 1125. https: // doi. org / 10.11609 / JoTT. ZPJ. 18.6.1117 - 25","Deuti, K & Ayyaswamy, A. K. (2008) Three new records of amphibians from West Bengal. Zoological Survey of India, 108 (4), 17 - 20.","Mathew, R. & Sen, N. (2010) Pictorial Guide to Amphibians of North East India. Zoological Survey of India, Kolkata, 144 pp.","Sangma, M. A. & Saikia, P. K. (2015) Checklist of amphibians inventoried from Tura Peak of West Garo Hills, Meghalaya, India. Journal of Global Biosciences, 4 (5), 2404 - 2424."]}

Megophrys (Xenophrys) zhangi Ye and Fei, 1992 (Figure 10) Megophrys zhangi Ye and Fei 1992:50 –52. In: A new pelobatid toad of the genus Megophrys from Xizang, China. Acta Herpetologica Sinica, 1–2: 50–52. Holotype. Adult male (CIB 750296: Figure 10), from “Zhangmo, Nyanang, Xizang altitude 1000 m ” (= Zhangmu town, ~ 27°59'24"N, 85°58'48"E, Nyalam County, Shigatse Prefecture, Tibet [or Xizang] Autonomous Region, China), collected by Yongzu Zhang on 26 June 1975 (Ye & Fei 1992). Paratypes. Two adult males (CIB 750295, CIB 750297), from the “same locality and date as the holotype, altitude 700–1000m ”, collector not mentioned (presumably the same). Examined specimen. Adult male (CIB 750296––photos only), holotype. Description of type series. Refer to Ye and Fei (1992) for the description in Chinese language text. Morphological comparison. Adult body size range provided for Megophrys zhangi is from the original description (Ye & Fei 1992). Megophrys zhangi (adult males, N =3) differs from M. monticola by its smaller adult body size, male SVL 32.5–37.2 mm (vs. adult male SVL 38.2–49.5 mm, N =17); differs from M. mangshanensis by absence of distinct white upper lip stripe (vs. present). For comparisons with subsequent species covered in this paper, refer to relevant morphological comparison sections for those species. Systematic position. Refer to the Systematic position section for Megophrys monticola above. Etymology. The specific epithet “ zhangi ” is a patronym, named for Yongzu Zhang, the collector of the type series. Suggested common name: Zhang’s Horned Frog. Distribution. This species is known with certainty only from the type specimens collected from between 700 and 1000 m elevation, below Zhangmu town (27°59′24″N, 85°58′48″E, 2300 m elevation), Nyalam County, Shigatse Prefecture, Tibet (or Xizang) Autonomous Region (Figure 8A). The proximity of the type locality to the Nepal border suggests that this species will also be found in adjacent areas of central Nepal., Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on page 30, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/2610202, {"references":["Fei, L., Ye, C. - Y. & Huang, Y. - Z. (1992) Two new species of the genus Megophrys, Pelobatidae (Amphibia, Anura) from China. Zoological Research, 13 (1), 5 - 12."]}

Mahony, Stephen, Teeling, Emma C., Biju, S. D. (2013): Three new species of horned frogs, Megophrys (Amphibia: Megophryidae), from northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region. Zootaxa 3722 (2): 143-169, DOI: http://dx.doi.org/10.11646/zootaxa.3722.2.2

Megophrys oropedion sp. nov. (Figures 3, 5��� 7; Table 1) ��� Megophrys monticola ���: Pillai and Chanda (1979: 386); Dutta (1997: 37 ���part; ���Shillong Peak���) ��� Megophrys parva ���: Chanda (1994: 19, 21���part; ���Shillong, Khasi Hills���); Chanda (1995: 459 ���part; ���Shillong���); Dutta (1997: 38 ���part; ���Shillong���); Chanda (2002: 11 ���part; ���Meghalaya���) ��� Xenophrys cf. parva ���: Mahony (2008: 9); Das et al. (2010: 54) Holotype. Adult male (ZSI A 11601), from Um Risa stream (25 �� 33.644 ���N 91 �� 52.960 ���E, 1,520 m asl.), Malki Forest (on Shillong Peak), Shillong, East Khasi Hills district, Meghalaya state, northeast India, collected by Systematics Lab members, 18 May 2009. Paratypes. Three adult males (ZSI A 11602, BNHS 5595, BNHS 5596), collected with holotype; adult female (ZSI A 11603), from Um Risa stream (25 �� 33.327 ���N 91 �� 53.099 ���E, 1,600 m asl.), Malki Forest, Shillong, East Khasi Hills district, Meghalaya state, northeast India, collected by Systematics Lab members, 22 May 2009. Referred specimens. Three adult males (SDB.DU 2009.300 ��� 301, SDB.DU 2009.1164) and one adult female (SDB.DU 2009.299), from Mawphlang Sacred Forest (25 �� 26.617 ���N 91 �� 44.767 ���E, 1,810 m asl.), Mawphlang, East Khasi Hills district, Meghalaya state, northeast India, collected by Systematics Lab members, 30 May 2009. Diagnosis. Megophrys oropedion sp. nov. is diagnosable from geographically relevant congeners by the following combination of characters: small-sized, robust species, adult-male SVL 32.8���39.2 mm (N= 7), adultfemale SVL 44.1���48.7 mm (N= 2); HW���HL, HW/SVL 33.9���38.5 % (N= 9); tympanum clearly defined, TYD /ED 53.7 ���69.0% (N= 9); SHL/SVL 41.7���48.4 % (N= 9), mean male SHL/SVL 45 % (N= 7), mean female SHL/SVL 42.1 % (N= 2); vomerine ridges ovoid, with vomerine teeth present; subarticular, inner and outer palmar tubercles and inner metatarsal tubercles absent in life and in preservation; digit tips not expanded relative to digit width; webbing and lateral fringes on toes absent; dorsolateral folds always present, approximately 75 percent of trunk length; distinct ���X���-shape marking on dorsum absent; protruding projection posterior to cloaca of males absent; single dominant frequency range of male advertisement call at 3,100���4,250 kHz, with maximum note energy of between 3,364���3,527 kHz (3,388 �� 70 kHz) at 18 ��C (Figure 7). Comparisons. Megophrys oropedion sp. nov. differs from the following large-sized species by considerably smaller adult-male size, SVL 32.8���39.2 mm, N= 7 (vs. adult-male SVL> 53 mm: M. gigantica, N= 6; M. glandulosa, N= 10; M. jingdongensis, N= 2; M. lekaguli, N= 8; M. major, N= 8; M. medogensis, N= 16; M. omeimontis, N= 10; M. robusta, N= 4; M. spinata, N= 2); from the following medium-sized species by smaller adultmale size, SVL 32.8���39.2 mm, N= 7 (vs. Megophrys ancrae sp. nov. male SVL 39.1 ���45.0 mm, N= 8; M. binlingensis SVL 45.1 ���51.0 mm, N= 3; M. megacephala SVL 45.9���53.4 mm, N= 7; M. takensis SVL 47.3 ���53.0 mm, N= 3), further from Megophrys ancrae sp. nov. by HW���HL (vs. HL>HW), digit tips not expanded terminally (vs. digit tips distinctly expanded terminally), small sharp horn-like tubercle absent on the eyelids (vs. present on most individuals), further from M. binlingensis by lateral fringes on toes absent (vs. present), further from M. megacephala by considerably narrower head HW/SVL 33.9���38.5 %, N= 9 (vs. HW/SVL 40.2���45.1 %, N= 9), and further from M. takensis by vomerine ridges ovoid (vs. elongated stalk-like projections); from the following smallsized species by larger adult size, male SVL 32.8���39.2 mm, N= 7, female SVL 44.1���48.7 mm, N= 2 (vs. M. wuliangshanensis male SVL 27.3���31.6 mm, N= 10, female SVL 41.0��� 41.5 mm, N= 2; M. zunhebotoensis male SVL 30.0 mm, N= 1, female SVL 39.0 mm, N= 1), further from M. wuliangshanensis by larger tympanum on males, mean TYD /ED 60.4 %, N= 7 (vs. mean male TYD /ED 48.8 %, N= 10), considerably shorter mean shank length, mean male SHL/SVL 45 %, N= 7, mean female SHL/SVL 42.1 %, N= 2 (vs. mean male SHL/SVL 50.5 %, N= 10, mean female SHL/SVL 50.4 %, N= 2), and further from M. zunhebotoensis by male SHL/SVL 43.4���48.4 %, N= 7, female SHL/SVL 41.7���42.6 %, N= 2 (vs. male SHL/SVL 50 %, N= 1, female SHL/SVL 46.1 %, N= 1); from M. nankiangensis and M. shapingensis by presence of clearly defined tympanum (vs. tympanum concealed by supratympanic fold); from M. binchuanensis and M. wushanensis by subarticular tubercles absent (vs. present), and lateral fringes on toes absent (vs. present); from M. minor by HW���HL (vs. HL>HW), vomerine teeth present (vs. absent), differences in male advertisement call characters, note duration 30���55 ms (44.6 �� 7.33 ms, N= 32) and a single primary frequency range at 18 ��C (vs. note duration 74.6���110.1 ms [90.8 �� 12.6 ms, N= 14], and a second subdominant frequency pulse at 6,800���7,600 kHz at 14 ��C); from M. palpebralespinosa by webbing between toes absent (vs. approx. half digit length); from M. daweimontis by inner and outer palmar tubercles and inner metatarsal tubercles absent in life and in preservation (vs. distinct and red in colour), SHL/SVL 41.7���48.4 %, N= 9 (vs. SHL/SVL 53.7 % on holotype); from M. wawuensis and M. zhangi by distinct ���X���-shape marking on dorsum absent (vs. present) and dorsolateral folds distinct (vs. indistinct), further from M. wawuensis by larger tympanum, TYD /ED 53.7 ���69.0%, N= 9 (vs. TYD /ED ca. 33 %), and further from M. zhangi by lateral fringes on toes absent (vs. present); from M. pachyproctus by protruding projection posterior to cloaca of male absent (vs. present); from M. parva by larger adult female size, SVL 44.1���48.7 mm, N= 2 (vs. female SVL 41.1���41.4 mm, N= 2), larger tympanum TYD /ED 53.7 ���69.0%, N= 9 (vs. TYD /ED 40.0��� 48.9 %, N= 4), and digit tips not expanded relative to digit width (vs. digit tips noticeably expanded relative to digit width); from M. serchhipii by dorsolateral folds always present, approximately 75 percent of trunk length (vs. absent). Holotype description (measurements in mm). Mature male (SVL 32.8) (Figure 5, 6C). Head small (HW 12.0, HL 12.0, IFE 6.0, IBE 10.1), as wide as long; snout rounded in dorsal view, obtusely protruding in lateral view, without rostral appendage (Figure 5 C); loreal region vertical and concave; canthus rostralis angular; dorsal region of snout slightly concave; eye (EL 4.4) longer than maximum tympanum diameter (TYD 2.7), and subequal to snout (SL 4.5); eye-tympanum distance (TYE 1.8) shorter than maximum tympanum diameter; tympanum slightly oval, vertically orientated, not concealed by supratympanic ridge (Figure 5 C); pupil in life oval, horizontally orientated when dilated and vertically when constricted; nostril orientated laterally, situated mid-way between eye and snout (EN 2.2, SN 2.3); internarial distance (IN 3.8) equal to eyelid width (UEW 3.8), and wider than the narrowest point between upper eyelids (IUE 3.4); pineal ocellus not visible externally; vomerine ridges moderately raised, ovoid, positioned at level posterior to choanae, separated from each other by distance equal to distance from choanae; vomerine teeth small; tongue moderately large, weakly notched posteriorly, with no medial lingual process. Forelimbs moderately short and thin, forearm (FAL 7.6) slightly enlarged relative to upper forelimb, shorter than hand (HAL 10.1); fingers short and rounded without lateral fringes (Figure 5 D), finger length formula I=II=IVColour in preservative (Figure 5): Entire dorsal and lateral surfaces of head, body, forelimbs and hindlimbs light brown; solid darker brown triangular marking between eyes, and thin dark brown stripe extending from tip of snout posterodorsally to centre of snout; supratympanic folds whitish-cream; front of snout with dark brown blotches; wide vertical dark brown bar below eyes and dark brown blotches covers tympanum; two dark brown blotches on anterior lateral surface of forearm, and dorsal surface of hands and feet with dark brown spots and speckles; feint dark brown transverse bands present on dorsal surfaces of hindlimbs. Gular region, chest, abdomen and ventral surface of limbs pale greyish-brown; feint, dark mottling on limbs; some small white spots along margin of lower mandible and distinct dark brown patch on either side of posterior ends of mandibles extending to insertion of forelimbs; ventral surfaces of tarsus and feet dark grey brown; area surrounding vent and posterior surfaces of thighs dark brown; ventral surface of hands grey-brown; pectoral and femoral glands white. Colour in life (Figure 6 C): As described above, except that dorsal colour was a richer mid-brown with tips of dorsal tubercles and large granules orange; inner surface of thighs and groin with some orange colouration. Iris colour metallic brown. Variation. See Table 1 for morphometric characters of seven adult males and two adult females. Finger length formula varies considerably in this species with the following combinations found in the series examined: I���II���IVSecondary sexual characters. Males have slightly raised nuptial pads covered with black micro-granules, covering most of the dorsal surface of the base of finger I, narrowing distally and extending to the base of the distal phalange on the inner dorsal side; nuptial pad medium sized and oval shaped on finger II, on the inner dorsal side of the base of the digit extending to the mid-proximal phalange; external vocal sac distinct as loose skin on most specimens, and forms a large single subgular sac that extends onto the anterior chest when fully inflated; internal vocal slit present on the floor of the mouth, near the rear of the lower mandible on both sides; forearm slightly to moderately enlarged relative to upper forelimb; protruding fleshy projection posterior to cloaca absent. Females lack nuptial pads, external vocal sac, internal vocal slits and enlarged forearms���ova are unpigmented. Etymology. The specific epithet ��� oropedion ��� is Greek meaning ���plateau��� or ���tableland���, as all known collection localities for the species are situated on the upper reaches of the Shillong Plateau. We treat the specific epithet as a noun in apposition to the generic name. Distribution. This species is confirmed here from Malki Forest (1,520���1,600 m asl.), Shillong, and Mawphlang Sacred Forest (1,810 m asl.), Mawphlang village, both in the East Khasi Hills district of Meghalaya state, northeast India (Figure 3). It has also been observed, but not collected from North East Hill University (NEHU) (25 �� 36.767 ���N 91 �� 45.050 ���E, 1,410 m asl.), on the outskirts of Shillong city (SM pers. obs.). This species is likely to be found in other forest fragments in the East and West Khasi Hills districts that cover the upper reaches of the Shillong Plateau above 1,400 m asl. Habitat and natural history. The forest type around Shillong and Mawphlang is categorized as subtropical wet hill forest (Champion & Seth 1968). The forest at the type locality is dominated by pine (Pinus kesiya Royle & Gordon) plantation, with a considerable amount of broadleaf trees lining the stream banks (SM, pers. obs.) (Figure 4 B), while the forest at Mawphlang is dominated by broadleaf trees (Figure 4 C). The collection streams at these localities were approximately 1���3 m wide and 10���40 cm deep, of moderate flow on a slight incline, primarily rocky with intermittent gravel bed, and with quiet pools that accumulate detritus, branches and leaf litter (for further details on habitat at Mawphlang see Mahony [2008]). The species was only found along portions of the streams that experience low levels of human disturbance. One female was collected on a trail ca. 200 m from the nearest stream, and another collected from a trail nearby a stream. Males were collected from the lower branches of bushy vegetation at ca. 20 cm above ground level, within 2 m from the edge of streams, or from rocks on the stream banks. One uncollected male was located inactive during day time in a fallen hollow branch lying over a stream (1 m above stream level). Metamorphosing juveniles were found during May and June at Shillong and Mawphlang. Males began calling vigorously from dusk until late into the evening (18:00 until at least 23: 30 h), regardless of precipitation level, but more intensely during light rain and/or when cloud cover envelops the forest, when regular calling began as early as 16:00 h, but was reduced during heavy precipitation. Calling was observed during visits between 20 th May to 15 th September. Sporadic daytime calling was heard on both overcast and clear days at Shillong and Mawphlang, and it consisted of one to two call repetitions followed by extended silence, usually from unobserved calling sites within dense bushes. Calling males distanced themselves from others by a minimum of 1 m of stream bank and distinctive territorial calls were emitted when males were played back their call at close quarters, as noted in other species (Mahony & Reza 2008). The advertisement call of Megoph, Published as part of Mahony, Stephen, Teeling, Emma C. & Biju, S. D., 2013, Three new species of horned frogs, Megophrys (Amphibia: Megophryidae), from northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region, pp. 143-169 in Zootaxa 3722 (2) on pages 151-157, DOI: 10.11646/zootaxa.3722.2.2, http://zenodo.org/record/249321

Megophrys vegrandis sp. nov. (Figures 3, 8 & 9; Table 1) “ Xenophrys boettgeri ?”: Athreya (2006: 100) “ Xenophrys cf. boettgeri ”: Athreya (2006: 139, 143) “ Xenophrys boettgeri ”: Mathew and Sen (2010: 54, fig. 89) Holotype. Adult male (ZSI A 11605), from a tributary of the Sessa Nadi (river), Sessa village (27 °06.067’N 92 ° 31.642 ’E, 1,110 m asl), Bhalukpong Forest Division, West Kameng district, Arunachal Pradesh state, northeast India, collected by Systematics Lab members, 8 August 2009. Paratypes by original designation. Two adult males (ZSI A 11604, BNHS 5597), collected with holotype. Referred specimen. One adult male (SDB.DU 2009.1274), and one metamorphosed juvenile (SDB.DU 2009.1275), collected with holotype. Diagnosis. Megophrys vegrandis sp. nov. is diagnosable from geographically relevant congeners by the following combination of characters: small-sized, slender species, adult male size amongst the smallest of all known Megophrys species, SVL 27.5–30.6 mm (N= 4); tympanum circular, not concealed by supratympanic ridge; vomerine ridges and vomerine teeth absent; dorsolateral folds extending for approximately 20–40 percent of trunk length; length of finger I 50 –65 percent of length of finger II; webbing between toes rudimentary; lateral fringes on toes present; nuptial pads, subarticular tubercles and protruding projection posterior to cloaca of males absent; groin and ventral thigh colouration not contrasting with surrounding regions on males; ventral surface of thighs mottled and blotched with dark brown. Comparisons. Megophrys vegrandis sp. nov. differs from the following large species (in parenthesis) by considerably smaller adult-male size, SVL 27.5–30.6 mm, N= 4 (vs. adult-male SVL> 53 mm: M. gigantica, N= 6; M. glandulosa, N= 10; M. jingdongensis, N= 2; M. lekaguli, N= 8; M. major, N= 8; M. medogensis, N= 16; M. omeimontis, N= 10; M. robusta, N= 4; M. spinata, N= 2); from the following small to medium-sized species (in parenthesis) by smaller adult-male size, SVL 27.5–30.6 mm, N= 4 (vs. Megophrys ancrae sp. nov. SVL 39.1 –45.0 mm, N= 8; M. binchuanensis SVL 32.0–36.0 mm, N= 4; M. binlingensis SVL 45.1 –51.0 mm, N= 3; M. daweimontis SVL 34–37 mm, N= 18; M. megacephala SVL 45.9–53.4 mm, N= 7; M. minor SVL 32.2–40.5 mm, N= 15; M. pachyproctus SVL 35.3–36.2 mm, N= 2; M. palpebralespinosa SVL 36.2 –38.0 mm, N= 2; M. parva SVL 33.9–41.4 mm, N= 4; Megophrys oropedion sp. nov. male SVL 32.8–39.2 mm, N= 7; M. takensis SVL 47.3 –53.0 mm, N= 3; M. wawuensis SVL 34.4–42.8 mm, N= 4; M. zhangi SVL 32.5–37.2 mm, N= 3), further from Megophrys ancrae sp. nov., M. megacephala, M. minor, Megophrys oropedion sp. nov., M. parva, and M. takensis by lateral fringes on toes present (vs. absent), further from M. binchuanensis by subarticular tubercles absent (vs. present), groin colouration not contrasting to surrounding regions on males (vs. red on males), further from M. binlingensis by vomerine ridges absent (vs. present, slender), further from M. daweimontis and M. wawuensis by FIL 50–65 % FIIL (vs. FIILM. pachyproctus by protruding projection posterior to cloaca of male absent (vs. present), FIL 50–65 % FIIL (vs. FIL subequal to FIIL), tympanum circular (vs. oval), and lateral fringes on toes present (vs. absent), further from M. palpebralespinosa by webbing between toes rudimentary (vs. approx. half digit length), further from M. zhangi by ventral thighs mottled and blotched with dark brown (vs. immaculate); from M. nankiangensis and M. shapingensis by presence of clearly defined tympanum (vs. tympanum concealed by supratympanic fold); from M. wushanensis by subarticular tubercles absent (vs. present), groin colouration not contrasting with surrounding regions on males (vs. red on males); from M. serchhipii and M. zunhebotoensis by lateral fringes on toes present (vs. absent), and vomerine teeth absent (vs. present); from M. wuliangshanensis by lateral fringes on toes present (vs. absent), dorsolateral folds extend approximately 20–40 percent body length (vs. most of body length), ventral thigh colouration not contrasting with surrounding regions on males (vs. red on males). Holotype description (measurements in mm). Mature male (SVL 29.2) (Figure 8, 9). Head small (HW 10.2, HL 10.2, IFE 5.2, IBE 9.0), as wide as long; snout rounded in dorsal view, obtusely protruding in lateral view, without rostral appendage (Figure 8 C); loreal region vertical and concave; canthus rostralis blunt; dorsal region of snout slightly concave (Figure 9 C); eye (EL 3.6) twice as long as maximum tympanum diameter (TYD 1.8), and shorter than snout (SL 4.0); eye-tympanum distance (TYE 1.5) shorter than maximum tympanum diameter; tympanum circular, with upper margin concealed by supratympanic ridge (Figure 8 C); pupil in life oval, horizontally orientated when dilated; nostril oriented laterally, situated mid-way between eye and snout (EN 2.0, SN 2.0); internarial distance (IN 3.3) subequal to eyelid width (UEW 3.3), and narrower than narrowest point between upper eyelids (IUE 3.7); pineal ocellus not visible externally; vomerine ridges and vomerine teeth absent; tongue moderately large, weakly notched posteriorly, with no medial lingual process. Forelimbs moderately long and thin, forearm (FAL 7.6) slightly enlarged relative to upper forelimb, shorter than hand (HAL 9.4); fingers short and broadly flattened but without lateral fringes (Figure 8 D), finger length formula I Skin of dorsal and ventral surfaces of head, body and limbs primarily smooth; flanks with small scattered tubercles (Figure 9 A); tympanum smooth, borders slightly raised, surrounding area posterior to eye granular (Figure 8 C); small tubercle present on outer edge of upper eyelid; supratympanic fold narrow without widening posteriorly, extends from orbit and curves down broadly through upper border of tympanum terminating above shoulder (Figure 8 C); short thin dorsolateral fold extending from behind supratympanic fold to approximately one third distance to groin; weak “V”-shaped parietoscapular ridge present, its two sides extending posteriorly from above tympanum and meeting medially beyond level of axilla; second inverted “V”-shaped ridge present on mid-dorsum (Figure 8 A); pectoral glands small and raised slightly, positioned on level with axilla (Figure 8 B); femoral glands small slightly raised, positioned sub-equally distant from knee and cloaca. Skin asperities absent on all surfaces. Colour in preservative (Figure 8): Dorsal and lateral surfaces of head and body primarily light grey-brown; brown triangular marking with light central blotch between eyes, and large brown, roughly “X”-shaped blotch present on dorsal surface of body; lower borders of dorsolateral and supratympanic folds, and flank tubercles dark brown; front of snout mottled with dark brown; wide oblique dark brown bar below eye and dark brown blotch covers tympanum; lateral surfaces of forearm and dorsal surface of hands and feet with dark brown spots and speckles; forelimbs brown above with feint dark brown transverse stripes on forearm; dorsal surface of upper forelimb pale relative to remaining dorsal trunk colour; dorsal surface of hindlimbs light grey-brown with oblique dark brown crossbars. Gular region, chest and anterior part of abdomen primarily greyish-yellow with barely discernible mottling and lighter patches along margin of lower mandibles; light colouration of chest darkens slightly posteriorly on abdomen which is mottled with light brown; broad dark brown row of blotches present on ventrolateral surfaces of abdomen; ventral surfaces of thighs and shanks with light and dark brown mottling; ventral surfaces of tarsus and feet dark grey-brown; area surrounding cloaca and posterior surfaces of thighs dark brown; ventral surface of hands grey-brown; pectoral and femoral glands lighter than surrounding area. Colour in life (Figure 9): Dorsum of head, body, hindlimbs and forearm light olive brown; above mentioned dorsal markings of head, body and limbs mid-brown; lateral surfaces of body and hindlimbs light mauve-grey; dorsal surfaces of upper forelimbs plain yellow; all dorsal and lateral tubercles, ridges and folds on head, body and limbs, orange; ventral surfaces of tips of fingers I and II and tips of all toes, orange. Iris metallic yellowish-orange. Variation. See Table 1 for morphometric characters of four adult males. Dorsal markings of paratypes agree with holotype, however overall colouration is darker. The length of the dorsolateral folds on paratypes and the adult referred specimen varies from approximately 20–40 percent trunk length. The supratympanic fold does not obscure the upper border of the tympanum on ZSI A 11604 and SDB.DU 2009.1274. Pineal body externally visible on ZSI A 11604 and BNHS 5597. BNHS 5597 has very sparse, small clear asperities on the dorsal folds of the body and hindlimbs and a few on the outer margin of the lower jaw, whereas SDB.DU 2009.1274 has sparse asperities on the posterior part of the dorsum only. ZSI A 11604, BNHS 5597, and SDB.DU 2009.1274 possess a short ridge on the upper eyelid as opposed to the tubercle present on the holotype. Dark blotches on the ventral surfaces of the body and limbs vary between specimens with respect to density and shade. The dorsal surface of the upper forelimbs vary between yellow and light brown in life. On the metamorph (SDB.DU 2009.1275), the dorsal and lateral surfaces of the head and body are mottled brown with faintly darker brown triangle between the eyes. Limbs greyish-yellow with feint brown crossbars. Lateral head markings like on the adults. Throat and abdomen pale grey with dark brown longitudinal stripe on each side of the lower flanks, on posterior thighs across the cloacal region and on the ventral tarsus. Pectoral and femoral glands visible as white spots. Secondary sexual characters. Males without nuptial pads on fingers; external vocal sac indistinct; internal vocal slits present near the rear of the lower mandible; protruding fleshy projection posterior to cloaca absent. Females currently unknown. Etymology. The specific epithet “ vegrandis ” is a Latin word meaning “diminutive” or “tiny” with reference to the small adult male size of this species. Distribution. The species is known from the type locality near Sessa village, in West Kameng district, in western Arunachal Pradesh state. We provisionally assign another population to this species from Sessni (27 °02.844’N 92 ° 25.086 ’E, 1,250 m asl), Eaglenest Wildlife Sanctuary, approximately 12.4 km west of the type locality (Athreya 2006: see remarks below). It is likely that the species is more widespread in suitable habitat at similar altitudes in western Arunachal Pradesh. Habitat and natural history. The primary habitat at the type locality is referred to as subtropical forest (Athreya 2006). The type specimens were collected from the banks of a shallow stream (max. depth 10 cm) that flowed through an area of dense bushy vegetation surrounded by disturbed, selectively logged forest (Figure 4 A). The vegetation formed a low complete canopy (1–2 m height) over the stream. The stream bed consisted of gravel and rocks, with leaf litter gathered in snags and side pools. No individuals of this species were found (or heard) from the banks of the larger Sessa Nadi river. All males were found calling from atop large boulders bordering the stream where individuals were spaced a minimum of three meters apart along the stream bank. Calling began at dusk on a dry cloudless night. The newly metamorphosed specimen was collected on the gravel bank of the stream. The vigor of male vocalisations indicates that the breeding season includes early August. Remarks. Athreya (2006) provided a selection of photographs, some basic measurements and observations of living individuals from Sessni, Eaglenest Wildlife Sanctuary which they identified as “ Xenophrys cf. boettgeri ” (pg. 143; and pg. 100, plate 2 as “ Xenophrys boettgeri ?”) and which appear to be conspecific with Megophrys vegrandis sp. nov. Mathew and Sen (2010: 54, fig. 89) also show a figure of a Megophrys vegrandis sp. nov. individual from Sessni, and provided a brief description as “ Xenophrys boettgeri ”, which they claim to be found in Arunachal Pradesh and Sikkim. Several of the characters they provide in the description (“tympanum...as large as eye”; “TTA [tibiotarsal articulation] reaches tip of snout”), however, neither match M. boettgeri s.s., or the photographed Megophrys vegrandis sp. nov, thus it is unclear what species they are basing their description. Morphologically, M. boettgeri and Megophrys vegrandis sp. nov. differ considerably (e.g., SVL of adult males 34.5–37.8 mm in M. boettgeri [vs. 27.5–30.6 mm in Megophrys vegrandis], lateral fringes on toes absent [vs. present], subarticular tubercles present on fingers [vs. absent], dorsolateral folds absent [vs. present anteriorly]) and thus these species can hardly be confused. Many other reports of M. boettgeri exist in the Indian literature primarily based on a misunderstanding that originated with Annandale (1917). Annandale regarded tadpoles of M. boettgeri (= M. brachykolos Inger & Romer—fide Inger & Romer 1961) from Hong Kong, to be superficially similar to tadpoles from Arunachal Pradesh which he had earlier assigned to Megalophrys kempii Annandale (Annandale 1912). He conjectured that Megalophrys kempii may therefore be synonymous with M. boettgeri, strangely disregarding the clear morphological differences between the adults of both species. Many subsequent authors listed M. boettgeri as present in Arunachal Pradesh based directly or indirectly on this weakly supported synonymy and not based on subsequent specimen collections (i.e., Bordoloi et al. 2000; Chanda 1990, 1992, 1994, 2002; Pillai & Chanda 1976; Sarkar & Ray 2006). Delorme et al. (2006) referred Megalophrys kempii to the rhacophorid genus Philautus Gistel based on a morphological examination of the holotype. The tadpoles that Annandale (1912, 1917) provisionally assigned to “ P.” kempii are therefore referable to an unknown species of Megophrys, thus rendering inaccurate all of the aforementioned publications that list M. boettgeri from Arunachal Pradesh. Dutta (1997) recognised “ P.” kempii (as Megophrys kempii) as valid, but unusually gives Sikkim as the range of M. boettgeri in India, and states that “After, Boulenger (1894) the species has not been recorded from India ”. The reference for this citation is not in the bibliography section, and here is regarded questionable as it predates the description of M. boettgeri (Boulenger 1899), and furthermore Sikkim is not included in the distribution of this species in Boulenger’s subsequent review of the group (Boulenger 1908). Pawar & Birand (2001) reported sighting M. cf. boettgeri from Mouling National Park in Arunachal Pradesh, from an unspecified source from Bhalpakram National Park in Meghalaya, as well as other localities in Meghalaya and Nagaland. Pawar & Birand (2001) recognised that further research was required to confirm the identity of the Indian populations of M. boettgeri. In the absence of referred voucher specimens, images, or any verifiable evidence of this species from their listed localities, we regard their report as anecdotal, and most likely misidentifications of another/other species. In summary, all literature reports of M. boettgeri from India were either based on Megophrys vegrandis sp. nov. (see above chresonymy), “ Philautus ” kempii (Bordoloi et al. 2000; Chanda 1990, 1992, 1994, 2002; Pillai & Chanda 1976; Sarkar & Ray 2006), or anecdotal records (Dutta 1997; Pawar & Birand 2001; Mathew & Sen 2010). No Indian voucher specimens referable to M. boettgeri have been specifically identified in literature, nor are any currently present in major museums, e.g., BMNH, ZSI, SDB.DU (Mahony pers. obs.). Outside India, the closest reported locality for M. boettgeri is from eastern Guangxi Province, China, ca. 1,400 km east of the eastern-most border of northeast India (Fei et al. 2009), thus in the absence of any solid evidence to support the presence of this species in India, it should no longer be included in the Indian amphibian checklists. The geographical distribution of M. boettgeri is here restricted to eastern China (Fei et al. 2009), where based on current understanding it should be considered endemic. (ind) (ind) (ind) (ind) (ind) (ind) (ind) (ind) (ind) (ind) (ind) (ind) 2.2 (ind) 2.2 (ind) 2.2 (ind) 2.5 (ind) (ind) (ind) (ind)

Megophrys ancrae sp. nov. (Figures 1���3; Table 1) Holotype. Adult male (ZSI A 11606), from 6 th mile (27 �� 29.833 ���N 96 �� 23.467 ���E, 420 m asl.) approx. 3 km from Deban Guest House, Deban, Namdapha National Park and Tiger Reserve, Changlang district, Arunachal Pradesh state, northeast India, collected by the Systematics Lab members, 2 June 2011. Paratypes. Adult male (ZSI A 11607), collected along with the holotype. Four adult males (BNHS 5598, BNHS 5600���5602), one subadult and one adult female (ZSI A 11608, BNHS 5599, respectively), from Bornulla forest stream (27 �� 32.367 ���N 96 �� 28.650 ���E, 790 m asl.), Namdapha National Park and Tiger Reserve, Changlang district, Arunachal Pradesh state, northeast India, collected by the Systematics Lab members, 3 May 2011. Referred specimens. Two adult males (SDB.DU 2009.727, 2009.730), from approx. 1 km before reaching Deban Guest House (27 �� 29.846 ���N 96 �� 23.470 ���E, 330 m asl.), Deban, Namdapha National Park and Tiger Reserve, Changlang district, Arunachal Pradesh state, northeast India, collected by the Systematics Lab members, 20 June 2009. Diagnosis. Megophrys ancrae sp. nov. is diagnosable from geographically relevant congeners by the following combination of characters: medium-sized slender species, adult-male SVL 39.1 ���45.0 mm (N= 8), adultfemale SVL 48.9 mm (N= 1); HL>HW, HW/SVL 30.0��� 35.8 % (N= 10); SHL/SVL 46.0��� 55.3 % (N= 10); tympanum clearly defined, TYD /ED 50.0��� 63.2 % (N= 10); vomerine ridges small to medium-sized, weakly raised and circular; distinct dorsolateral folds present; subarticular, palmar and metatarsal tubercles absent; lateral fringes on toes absent; webbing between toes rudimentary; digit tips distinctly expanded terminally; protruding projection posterior to cloaca of males absent; dark spots on the throat present on most specimens; small, sharp, horn-like tubercle present on the eyelids of most individuals. Comparisons. Megophrys ancrae sp. nov. differs from the following large-sized species by considerably smaller adult male size, SVL 39.1 ���45.0 mm, N= 8 (vs. adult male SVL> 53 mm: M. gigantica Liu, Hu & Yang, N = 6; M. glandulosa Fei, Ye & Huang, N = 10; M. jingdongensis Fei & Ye, N= 2; M. lekaguli Stuart, Chuaynkern, Chan-ard & Inger, N= 8; M. major (Boulenger), N= 8; M. medogensis Fei & Ye, N= 16; M. omeimontis Liu, N = 10; M. robusta (Boulenger), N= 4; M. spinata Liu & Hu, N= 2); from the following medium-sized species by smaller adult size, male SVL 39.1 ���45.0 mm, N= 8, female SVL 48.9 mm, N= 1 (vs. M. megacephala Mahony, Sengupta, Kamei & Biju male SVL 45.9���53.4 mm, N= 7, female SVL 64.4 mm, N= 1; M. takensis Mahony male SVL 47.3 ��� 53.0 mm, N= 3, female SVL 72.9 mm, N= 1), further from M. megacephala by considerably narrower head, HW/ SVL 30.0��� 35.8 %, N= 10 (vs. HW/SVL 40.2���45.1 %, N= 9), and further from M. takensis by vomerine ridges small to medium-sized, weakly raised and circular (vs. elongated stalk-like projections); from the following small-sized species by larger adult size, male SVL 39.1 ���45.0 mm, N= 8, female SVL 48.9 mm, N= 1 (vs. M. daweimontis Rao & Yang male SVL 34���37 mm, N= 17, female SVL 40���46 mm, N= 3; M. pachyproctus Huang male SVL 35.3���36.2 mm, N= 2, female unknown; M. parva male SVL 33.9 ���36.0 mm, N= 2, female SVL 41.1���41.4 mm, N= 2; M. wuliangshanensis Ye & Fei male SVL 27.3���31.6 mm, N= 10, female SVL 41.0��� 41.5 mm, N= 2; M. zunhebotoensis Mathew & Sen male SVL 30.0 mm, N= 1, female SVL 39.0 mm, N= 1), further from M. daweimontis by HL>HW (vs. HW���HL), palmar and metatarsal tubercles absent in life and in preservation (vs. inner and outer palmar tubercles and inner metatarsal tubercles distinct and red in life), further from M. pachyproctus by protruding projection posterior to cloaca of male absent (vs. present), and further from M. parva by TYD /ED 49.2���63.2 %, N= 10 (vs. TYD /ED 40���48.9 %, N= 4), shanks considerably longer on males, SHL/SVL 46���55.3 %, N= 8 (vs. male SHL/SVL 42.8���43.7 %, N= 2); from M. nankiangensis Liu & Hu and M. shapingensis Liu by tympanum clearly defined (vs. tympanum concealed by supratympanic fold); from M. binchuanensis Ye & Fei and M. wushanensis Ye & Fei by subarticular tubercles absent (vs. present), and lateral fringes on toes absent (vs. present); from M. palpebralespinosa Bourret by webbing between toes rudimentary (vs. approx. half digit length); from M. binlingensis Jiang, Fei & Ye and M. minor by lateral fringes on toes absent (vs. present), further from M. binlingensis by vomerine ridges weakly raised and circular (vs. slender and not dilated at its posterior region), and further from M. minor by dark spots on the throat present on most specimens (vs. absent); from M. wawuensis Fei, Jiang & Zheng and M. zhangi Ye & Fei by dorsolateral folds distinct (vs. indistinct), further from M. wawuensis by larger tympanum, TYD /ED 49.2���63.2 %, N= 10 (vs. TYD /ED ca. 33 %), and further from M. zhangi by lateral fringes on toes absent (vs. present); from M. serchhipii Mathew & Sen by dorsolateral folds always present, usually 2 / 3 to complete body length (vs. absent). Holotype description (measurements in mm). Mature male (SVL 45.3) (Figures 1, 2A). Head small (HW 14.6, HL 15.6, IFE 7.5, IBE 12.6), longer than wide; snout rounded in dorsal view, obtusely protruding in lateral view, without rostral appendage (Figure 1 C); loreal region vertical and concave; canthus rostralis angular; dorsal region of snout slightly concave; eye (EL 5.4) twice as long as maximum tympanum diameter (TYD 2.7) and subequal to snout (SL 5.5); eye-tympanum distance (TYE 2.7) equal to maximum tympanum diameter; tympanum slightly oval, orientated vertically, its upper approx. 5 percent concealed by supratympanic ridge (Figure 1 C); pupil in life oval, horizontally orientated when dilated; nostril orientated laterally, closer to eye than snout (EN 2.5, SN 3.4); internarial distance (IN 4.7) subequal to eyelid width (UEW 4.6), and greater than narrowest point between upper eyelids (IUE 4.2); pineal ocellus not visible externally; vomerine ridges present, circular and weakly raised with small vomerine teeth, positioned between choanae, separated from each other by distance equal to distance from choanae; tongue moderately large, weakly notched posteriorly, with no medial lingual process. Forelimbs moderately long and thin, forearm (FAL 10.3) slightly enlarged relative to upper forelimb, and shorter than hand (HAL 13.1); fingers long and narrow without lateral fringes (Figure 1 D), finger length formula IColour in preservative (Figure 1): Entire dorsal and lateral surfaces of head, body, forelimbs and hindlimbs brown; slightly darker brown triangular marking with light central blotch between eyes; dorsolateral and supratympanic folds and flank tubercles brownish-cream; front of snout and lateral canthus rostralis dark brown; wide vertical dark brown bar below eyes and dark brown blotch covering tympanum; two dark brown blotches on anterior lateral surface of forearms; dorsal surface of hands and feet with dark brown spots and speckles. Gular region, chest and anterior part of abdomen primarily dark brown, with white speckling along outer margin of ventral mandibles; dark colouration of chest lightening posteriorly on abdomen which is mottled light and dark brown; ventral surfaces of thighs and shanks with pale brown mottling; ventral surfaces of tarsus and feet dark grey brown; area surrounding vent and posterior surfaces of thighs dark brown; forelimbs ventrally mottled light and dark brown; grey-brown colouration of ventral side of hands extending as a blotch onto ventral surface of forearms; pectoral and femoral glands white. Colour in life (Figure 2 A): Dorsally light greyish-brown; dorsal and ventral surfaces with orange speckling, most dense on groin region, on flank tubercles and granules of dorsum, and most noticeable on ventral surfaces where it contrasts with dark grey colouration; ventral surface of throat with feint, dark longitudinal stripes. Iris metallic mid-brown. Variation. See Table 1 for morphometric characters of eight adult males and two females. Vomerine ridges small to medium-sized. Of the ten examined specimens only the holotype and the two referred specimens (SDB.DU 2009.727 and SDB.DU 2009.730) possess visible, small vomerine teeth. Dorsolateral folds extend approximately 65���100 percent of the body length except on the holotype which possesses one side that extends only ca. 50 percent body length. All specimens have a prominent pointed tubercle on the eyelid except BNHS 5601 on which tubercles are absent, and BNHS 5600 which possesses a tubercle on one side only. Most specimens possess a ���V���-shaped parietoscapular ridge and a second inverted ���V���-shaped ridge on the mid-dorsum which joins laterally with the dorsolateral folds, except BNHS 5600 and BNHS 5599 which do not possess the mid-dorsal ridge. The presence of dermal asperities on SDB.DU 2009.730, BNHS 5598, and BNHS 5601���5602 is similar to the state observed on the holotype. Dermal asperities on ZSI A 11607, like on the aforementioned specimens but considerably denser. On SDB.DU 2009.727 dorsal asperities are restricted to the posterior half to the back, upper flanks, and tympanic region; on BNHS 5600 asperities on the lower jaw are absent and only sparsely cover the posterior back and hindlimbs. The dark ventral regions on the holotype are lighter grey-brown on some individuals (e.g., ZSI A 11607, SDB.DU 2009.727, SDB.DU 2009.730). BNHS 5598���5602 and ZSI A 11608 have distinct dark longitudinal stripes on the throat, and feint crossbars dorsally on hindlimbs. BNHS 5598���5602, and SDB.DU 2009.730 have barely distinguishable dark brown blotches surrounding the ���V��� shaped dorsal folds; remaining specimens without distinct dorsal markings. Dorsal colouration in life varies from light to dark brown, with varying density of orange speckling which never predominates the overall colouration (Figure 2 B���C). Secondary sexual characters. Males with slightly raised nuptial pads covered with black micro-granules, covering most of the dorsal surface of the base of finger I, narrowing distally and extending onto the base of the distal phalange on the inner dorsal side; nuptial pad small sized and oval shaped on finger II, only on the inner dorsal side of the base of the digit; external vocal sac distinct on some individuals, and indistinct on others; internal vocal slits present near the rear of the lower mandible; forearms slightly enlarged relative to upper forelimbs; extensive dermal asperities present; protruding fleshy projection posterior to cloaca absent. Female (BNHS 5599) with large unpigmented ova (diameter 1.4���1.7 mm, N= 4); both examined females do not possess nuptial pads, enlarged forearms, vocal sac or slits, or any dermal asperities. Etymology. The specific epithet ��� ancrae ��� is Latin for ���of the valley��� as all collection localities for the type series were along the foothills of the Noa-Dihing river valley in Namdapha National Park and Tiger Reserve. Distribution. This species is thus far known only from the type locality and surrounding foothill areas (330��� 790 m asl.) in Namdapha National Park and Tiger Reserve in Changlang district of eastern Arunachal Pradesh, northeast India (Figure 3). Habitat and natural history. The primary habitat type in Namdapha National Park and Tiger Reserve is broadly regarded as tropical lowland evergreen forest (see Proctor et al. 1998, and references therein for further description of forest composition). Megophrys ancrae sp. nov. was found to be relatively abundant at the type locality, however they were quite difficult to locate. ZSI A 11606���11607 were collected from leaf litter on the forest floor. BNHS 5598 was collected from a bush overhanging the Bornulla stream at ca. 3 m above stream level (Figure 4 D). All specimens were active and collected between 18: 30 ��� 19: 30 h. The adult female BNHS 5599 was gravid and all males were calling when collected, indicating that the breeding season for this species extends at least between 20 th May to 20 th July. Bornulla stream is a steep rocky torrent that flows through dense forest habitat with dense undergrowth vegetation. Remarks. In the past couple of decades the results of only a few herpetological surveys in Namdapha National Park and Tiger Reserve have been published (e.g., Pawar & Birand 2001; Sarkar & Ray 2006; Sarkar & Sanyal 1985). From the Megophryinae subfamily, Pawar and Birand (2001) reported the species Megophrys robusta; a large bodied species which can not be confused with Megophrys ancrae sp. nov. Though these authors mentioned that some specimens from their northeast Indian survey were collected for later identification, they do not provide specimen details, thus their locality reports must be considered anecdotal. Sarkar & Ray (2006) report a single specimen which they identify as Megophrys major, with a brief description stating characters that clearly distinguish it from the species described here, i.e., SVL 67 mm and toes ���more or less one fourth webbed���. A single adult female Megophrys specimen (BMNH 1934.10.2.10) was collected by Captain Kingdon Ward from the Lohit Valley, Sadiya Frontier Tract, and discussed by Smith (1935) under the identification of M. minor. The Lohit River flows through the Lohit and Anjaw districts of Arunachal Pradesh and forms the adjacent major drainage system to the north of that created by the Noa-Dihing river valley. The condition of the specimen is quite dehydrated making determination of soft characters unreliable, however it is not considered to be conspecific with Megophrys ancrae sp. nov. due to small adult female size (SVL 34.5 mm), absence of vomerine ridges, and circular tympanum (Mahony, pers. obs.). Further fresh collections of the Lohit population are necessary to revise Smith���s identification, and thus the verification of M. minor in India., Published as part of Mahony, Stephen, Teeling, Emma C. & Biju, S. D., 2013, Three new species of horned frogs, Megophrys (Amphibia: Megophryidae), from northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region, pp. 143-169 in Zootaxa 3722 (2) on pages 145-151, DOI: 10.11646/zootaxa.3722.2.2, http://zenodo.org/record/249321