identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03D6878AFFD1022FFF73F894FE16FE39.text	03D6878AFFD1022FFF73F894FE16FE39.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megophrys (Xenophrys) monticola (Gunther 1864)	<div><p>Megophrys (Xenophrys) monticola (Günther, 1864)</p><p>(Figures 6 &amp; 7; Table 1)</p><p>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.</p><p>[?] 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.</p><p>[?] 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.</p><p>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).</p><p>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 &amp; H; SDBDU 2011.1048: Figure 7H) and one adult female (SDBDU 2011.1049: Figure 7F &amp; 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 &amp; G), from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=88.16627&amp;materialsCitation.latitude=26.997234" title="Search Plazi for locations around (long 88.16627/lat 26.997234)">Sukhiapokhri-Manebhanjan</a> 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 <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=88.3&amp;materialsCitation.latitude=26.945555" title="Search Plazi for locations around (long 88.3/lat 26.945555)">Bagora Range</a> 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 &amp; SDBDU 2011.420), from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=88.398056&amp;materialsCitation.latitude=26.88861" title="Search Plazi for locations around (long 88.398056/lat 26.88861)">Ghoramara area</a> (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 <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=88.39777&amp;materialsCitation.latitude=26.909445" title="Search Plazi for locations around (long 88.39777/lat 26.909445)">Stream</a> 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 <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=88.38402&amp;materialsCitation.latitude=26.884583" title="Search Plazi for locations around (long 88.38402/lat 26.884583)">Mahananda Wildlife Sanctuary</a> (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 <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=88.39848&amp;materialsCitation.latitude=26.9095" title="Search Plazi for locations around (long 88.39848/lat 26.9095)">Latpanchar town</a> (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 <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=88.3&amp;materialsCitation.latitude=26.945555" title="Search Plazi for locations around (long 88.3/lat 26.945555)">Makaibari Tea Estate</a> (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.</p><p>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.</p><p>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.</p><p>Forelimbs moderately long, thin (Figure 6A &amp; 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 I&lt;II&lt;IV&lt;III (FIL 3.6, FIIL 4.0, FIIIL 6.8, FIVL 4.2); interdigital webbing, lateral fringes, subarticular and supernumerary tubercles absent; thenar tubercle indistinct, metacarpal tubercles absent; finger tips appear slightly expanded, flattened without pads or terminal grooves. Hindlimbs relatively long, thin (Figure 6A &amp; B); thighs shorter than shanks and feet (TL 17.9, SHL 20.4, FOL 19.5); toes long, thin (Figure 6E), relative toe lengths I&lt;II&lt;V&lt;III&lt;IV; digit tips appear slightly dilated, flattened, without pads; webbing appears rudimentary; lateral fringes on toes indistinct; outer metatarsal, subarticular and supernumerary tubercles all absent; inner metatarsal tubercle indistinct; ridge of thickened skin absent on ventral surfaces of digits.</p><p>Skin texture is not described in detail due to the high level of dehydration (Figure 6), however, generally: dorsal surfaces of body and head (including upper eyelids) appear covered in fine granules; dorsolateral ridges composed of row of fine granules visible on anterior half of body, does not connect with supratympanic ridge on either side; weak V-shaped parietoscapular ridge present, extending posteriorly from temporal region, meeting medially beyond level of forelimbs; supratympanic ridges narrow with minimal widening posteriorly, extends from posterior orbital borders, curves down broadly through upper tympanum, terminating above forelimbs; ventral surfaces and limbs appear primarily smooth; pectoral and femoral glands not distinctly raised, pectoral glands positioned on chest level with axilla, femoral gland positioned on rear of each thigh at midpoint between knee and cloacal opening.</p><p>Colouration: In preservative (Figure 6A &amp; 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 &amp; E); pectoral and femoral glands lighter than surrounding surfaces. In life: Colouration not originally documented.</p><p>Description of referred specimen SDBDU 2011.1047 (measurements in mm). Mature male (SVL 40.7) (Figure 7C, E &amp; 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.</p><p>Forelimbs moderately long, thin (Figure 7C &amp; 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&lt;IV&lt;III (FIL 4.6, FIIL 4.6, FIIIL 8.1, FIVL 4.9); interdigital webbing, subarticular and supernumerary tubercles absent; outer metacarpal and thenar tubercles weakly developed; finger tips rounded, not expanded relative to digit widths, with poorly defined circular pads, terminal grooves on pads absent. Hindlimbs moderately long, thin (Figure 7C &amp; E); thighs as long as feet, and shorter than shanks (TL 17.9, FOL 17.9, SHL 18.9); toes long, without lateral fringes, relative toe lengths I&lt;II&lt;V=III&lt;IV; toe tips rounded, not dilated, with weakly defined longitudinally oval-shaped pads, terminal grooves on pads absent; inner metatarsal tubercle weak, without distinct borders; webbing, subarticular, supernumerary and outer metatarsal tubercles absent; callous tissue absent on ventral surface of all digits.</p><p>Skin of dorsal surfaces of body, upper forelimbs, dorsal and lateral surfaces of head weakly granular (Figure 7C); tympanum smooth with borders slightly raised; outer edge of upper eyelids with a short transverse ridge; supratympanic ridges narrow anteriorly gradually expanding posterior to tympanum to become moderately enlarged and glandular, extending obliquely from orbits, abruptly curving above posterior upper border of tympanum, terminating above forelimb insertions; flanks with small to large scattered pustular tubercles; dorsolateral ridges thin, well defined, extending posteriorly from behind supratympanic ridges to ~90% trunk length, non-continuous with gap on anterior third of its length; parietoscapular and sacral ridges weak, not connected, configuration as “&gt; &lt;”; short, obliquely transverse ridges present on dorsal surface of thighs, shanks and forearms; gular region, chest, abdomen and ventral surfaces of limbs smooth; pectoral glands small, weakly raised, level with axilla on chest; femoral glands moderately large, slightly raised, on posterior surface of thighs, subequally distant from knees and cloaca; medium sized white dermal asperities present, forming dense narrow circummarginal band on lower jaw, tympanic region (excluding tympanum), below and including supratympanic ridges and dorsal ridges, sparse on upper eyelids, posterior surfaces of head and anterior dorsum, increasing in density posteriorly to above (but not surrounding regions of) cloaca, few also on tibia, absent from all other surfaces.</p><p>Colouration: In preservative (Figure 7E &amp; 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.</p><p>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&lt;II&lt;IV&lt;III, I=II&lt;IV&lt;III, I=II=IV&lt;III, IV&lt;I=II&lt;III; occasionally toe III=V in length; vomerine ridge varies from small to medium sized, round to ovoid, positioned between to slightly posterior to level of choanae, occasionally appear to be slightly closer to choanae than to each other; small vomerine teeth present on most specimens; posterior edge of tongue weakly notched on most specimens, but appears rounded on others (likely artefact of preservation); dorsolateral ridge length varies from ~50 to 95% trunk length, can be prominently raised on some individuals; supratympanic ridges on most specimens broadly curve through upper portion of tympanum, concealing up to ~30% of tympanum; following parietoscapular-sacral ridge configurations were observed: “&gt; &lt;”, “&gt; (”, “&gt;–&lt;”, “&gt;–|”, “&gt;- &lt;”; dermal asperities coverage on males similar to that on referred specimen, typically varying in density of coverage on some surfaces, however asperities also found sparsely covering dorsal surfaces of thighs, tarsi, forelimbs, and on surrounding regions of cloaca; females have considerably less dermal asperities, juveniles have none; outer metacarpal and thenar tubercles moderately well developed on some individuals, weak inner metacarpal tubercle visible on some specimens; finger and toe tips very slightly expanded on some specimens. Distinct X-shaped dorsal marking present on bodies of approximately half of specimens; dorsal colouration varies between light yellowish-brown, orange-brown (Figure 7B), greyish-brown, or olive green (Figure 7C); groin and ventral surface of hindlimbs on many males and one female light to deep reddish-orange in life (Figure 7D); ventral markings vary considerably with small to large, dense to sparse blotching on abdomen (Figure 7D, G &amp; H), ventral surfaces of hindlimbs may be plain (without markings) or densely mottled with dark brown blotches, throat may be pale brown with distinct large darker blotches, or almost plain dark brown.</p><p>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 &lt;3 mm); nuptial pads, vocal sac, internal vocal slits, enlarged forearms, all absent.</p><p>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.</p><p>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 &amp; 4). In Darjeeling, this species has two comparatively deeply divergent mitochondrial haplotypes (Appendix I, Table 6; Appendix II, Figures 1, 5, &amp; 7). However, nuclear diversity observed between these populations consisted of pSNPs only (Figures 3 &amp; 5; Appendix II, Figures 3, 4, &amp; 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 current data does not suggest that the distribution of M. zhangi s.s. extends into India.</p><p>Etymology. The specific epithet “ monticola ” is a compound Latin word derived from montis meaning “mountains” and - cola meaning “dwelling in”.</p><p>Suggested common name: Mountain Horned Frog (Ahmed et al. 2009).</p><p>No. Status Sex SVL HW HL IFE IBE EL TYD TYE SL EN SN IUE IN UEW FAL HAL FIL FIIL FIIIL FIVL SHL TL TFOL FOL IMT</p><p>†1947.2. 25.19 LT F 102.0 42.0 40.9 16.9 30.1 11.6 7.7 8.2 13.2 5.3 7.4 12.0 11.9 10.0 25.4 26.3 12.5 12.5 18.2 12.2 52.6 50.0 73.6 50.8 7.4 ZSIC 9681 PLT F 108.3 42.8 40.1 - - 10.0 5.2 11.4 13.8 5.5 6.3 13.5 14.5 7.5 25.1 27.0 - - - - 54.6 53.5 73.0 - ind ZSIC 10777 PLT F 92.9 40.4 38.8 - - 9.7 7.2 9.9 13.3 6.5 6.6 13.2 11.8 8.4 23.1 25.0 - - - - 50.4 49.8 67.4 47.4 ind #2011.1063 RS F 81.3 31.3 30.1 14.9 24.0 9.1 3.6 7.3 11.2 4.9 6.1 10.6 9.5 7.8 18.2 22.1 10.4 10.4 14.9 9.4 42.4 36.0 57.2 38.1 4.4 #2011.416 RS M 83.1 32.7 31.5 15.7 25.7 10.5 4.0 9.0 11.4 5.2 6.6 9.7 10.0 9.1 19.5 21.9 10.1 10.1 14.1 9.9 42.2 39.6 57.0 37.0 5.2 #2011.1057 RS M 80.8 32.4 31.7 15.7 24.5 9.5 4.7 6.6 10.9 5.4 6.4 10.0 10.4 8.1 19.2 22.3 9.5 9.5 14.1 8.5 42.4 40.8 58.0 38.4 5.2 #2011.1062 RS M 79.8 32.7 31.3 15.6 23.9 10.4 4.6 6.4 11.2 5.0 6.4 10.1 9.5 8.1 18.8 21.7 10.6 10.6 14.7 10.2 42.6 39.4 59.6 39.4 5.1 #2011.1064 RS M 79.3 31.9 31.9 14.3 24.6 10.8 5.0 6.6 11.1 4.9 6.5 9.3 10.1 8.5 17.6 20.5 9.7 9.6 14.2 9.4 39.8 36.8 53.6 36.3 ind #2009.1270 RS M 81.2 29.7 31.2 14.4 23.0 8.9 4.6 5.7 11.4 5.2 6.1 9.4 9.4 7.1 16.7 22.2 9.4 9.4 13.9 9.4 42.2 39.1 58.6 40.3 5.2</p><p>……continued on the next page</p><p>BNHS 6041 PT F 68.0 25.8 26.3 11.8 19.2 8.1 4.1 6.1 9.1 3.6 5.4 6.6 7.5 6.7 14.1 17.0 8.4 8.1 11.4 7.9 36.9 33.0 50.0 33.5 ind BNHS 6042 PT F 68.0 27.4 28.2 11.8 19.4 8.1 3.8 6.1 9.0 3.6 5.4 6.4 8.3 6.5 15.5 17.4 8.6 8.6 11.7 7.3 39.2 35.7 51.4 34.7 ind BNHS 6043 PT F 74.6 29.4 28.5 11.9 21.4 8.1 3.7 6.3 9.0 3.9 5.8 7.0 8.7 7.8 16.2 18.2 8.7 8.7 12.1 7.9 41.8 39.4 54.2 37.9 ind BNHS 6040 HT M 68.4 26.3 25.0 10.5 18.1 7.6 3.1 5.8 8.6 3.8 5.4 5.4 7.7 6.8 14.6 17.9 8.7 8.6 11.9 7.8 36.7 35.5 51.0 34.7 ind #2009.297 RS M 56.9 23.5 23.7 10.0 17.0 7.2 3.7 4.4 7.5 3.1 4.6 5.7 7.4 6.0 13.9 15.1 7.4 7.4 10.6 6.8 33.9 30.8 45.8 29.1 ind BNHS 6044 PT M 62.4 23.2 23.3 10.4 17.2 7.6 3.9 5.1 7.9 3.8 5.0 5.4 7.2 6.1 14.2 16.8 7.8 7.4 11.0 7.6 34.4 31.4 47.7 31.6 ind #2007.134 RS M 67.5 25.2 25.2 10.8 18.4 8.8 4.1 4.9 8.9 3.6 5.2 6.2 7.9 6.2 15.2 18.2 8.9 8.8 12.2 8.5 41.1 36.9 57.2 37.9 ind #2009.298 RS J 33.8 13.7 13.6 6.2 10.5 4.5 2.0 2.8 5.0 2.4 2.9 3.4 4.1 3.9 7.2 8.0 3.6 3.6 5.0 3.1 16.5 15.1 23.1 15.2 ind</p><p>……continued on the next page</p><p>Distribution. Recent collections of this species were from mid to high-elevation areas (880–2220 m asl.). In the Darjeeling sub-division of Darjeeling district, it was found in Senchal Wildlife Sanctuary, and along the Sukhiapokhri-Manebhanjan road (Rambi range). In the Kurseong sub-division of Darjeeling district, it was found within the Bagora Range, Makaibari Tea Estate, Mahananda Wildlife Sanctuary, and countryside surrounding Latpanchar town. We also include localities provided for Megophrys sanu comb. nov. and Megophrys katabhako comb. nov. which are verifiable by molecular data from Deuti et al. (2017) in the Darjeeling district and neighbouring areas of Sikkim (Figure 8B) in anticipation of the conclusions of our taxonomic clarification of these taxa (see Remarks section below). Megophrys monticola appeared to be locally abundant at collection localities and presumably is more widespread in the Darjeeling Hills and neighbouring Sikkim state (e.g., Chanda 1986; Deuti et al. 2017; Subba et al. 2016 [as M. parva]). This species (as “ parva ”) has been widely reported from Nepal (Nanhoe &amp; Ouboter 1987; Anders 2002), although at least some of these populations in central and western Nepal may apply to M. zhangi, or similar unnamed species. This species is also likely to range east, at least into Bhutan; a photographed specimen in Wangyal and Gurung (2012) (as X. nankiangensis Liu &amp; Hu, 1966 [Hu et al. 1966]) appears to be superficially similar to the specimens examined in this study.</p><p>Habitat and natural history. Our observations of this species correspond well with details provided by Anders (2002) for the Nepalese populations. Most males in the SDBDU collection were found calling during May and June, along the banks of small temporary and permanent streams (&lt;30 cm to 2 m wide; e.g., Figure 9C) bordered by dense low vegetation. Here, males typically called from concealed positions on the ground, or from the lower branches of vegetation on the banks. Calling males typically separate themselves from their nearest competitor by at least two meters. Some specimens were collected from roadside streams where the adjacent bank consisted of a near vertical rocky wall, up to 2 m height above stream level, covered with low vegetation and mosses, where males concealed themselves in rocky crags. Females collected in early June contained large ova, indicating that this species breeds at least during the early monsoon season in Northeast India. No tadpoles were observed attributable to this species. Habitats surrounding the collection streams varied from mature primary growth forest to heavily disturbed secondary growth. Refer to Deuti et al. (2017) for further observations.</p><p>Remarks. Günther (1864) briefly described Xenophrys monticola as the sole member of the genus Xenophrys Günther, 1864 . The syntype series of X. monticola consisted of two specimens, one from “Sikkim” (a region now encompassing Sikkim state, and Darjeeling and Kalimpong districts of West Bengal state), and the other from “Khasya” (now the Khasi Hills, Meghalaya state). Examination of the two syntypes revealed that they represented two different species (Mahony et al. 2017). The accompanying figure in Günther’s (1864) original description clearly depicted the Sikkim specimen (Günther 1864: Pl. XXVI, fig. H; BMNH 1947.2.25.13). In order to provide nomenclatural stability to the genus-level name Xenophrys, restricting the name of the type species to a single biological species was warranted. Mahony et al. (2017) subsequently designated the Sikkim specimen as the lectotype of Xenophrys monticola Günther. The “Khasya” paralectotype (BMNH [18]53.8.12.52) has been provisionally considered to be synonymous with M. parva of eastern Myanmar by Deuti et al. (2017); however, we consider this specimen to be morphologically most similar to the Khasi Hills endemic, Megophrys oropedion Mahony, Teeling and Biju, 2013 .</p><p>After its description, Xenophrys monticola was reported from many localities in Northeast India, Nepal, Myanmar, and China (e.g., Boulenger 1882, 1890, 1893, 1899, 1907; Günther 1868; Sclater 1892b). However, its historically recognised extensive distribution is now understood to have resulted from misidentifications of other currently named and unnamed species. Boulenger (1893) described a small Megophrys species from eastern Myanmar as Leptobrachium parvum comparing it with specimens of M. major s.l., which he erroneously considered to represent X. monticola . Boulenger (1908) subsequently placed all megophryids in the genus Megalophrys . He recognised his previous error (noted in his chresonymy of M. major), and considered X. monticola to be conspecific with M. parva (= L. parvum), incorrectly retaining M. parva as the valid name for the taxa. Boulenger (1908) might have been led to this nomenclatural decision due to uncertainty regarding the valid name for the species Megophrys montana Kuhl and Van Hasselt, 1822a (also spelled Megalophrys monticola Kuhl &amp; Van Hasselt, 1822b) that he may have regarded to have been a senior homonym (see Dubois 1982, 1989, 1992 for discussions). Boulenger’s (1908) nomenclatural action went unquestioned and most subsequent reports of X. monticola s.s. in literature were under the name M. parva (e.g., Anders 2002; Boulenger 1908; Bourret 1942; Chanda 1986, 2002; Dutta 1997; Nanhoe &amp; Ouboter 1987; Sarkar et al. 1992; Subba et al. 2016, and many others). Subsequent reports of M. montana Kuhl and Van Hasselt (or M. monticola Kuhl &amp; Van Hasselt) from Northeast India (e.g., Annandale 1906; Pillai &amp; Chanda 1976, 1979, 1981; Dutta 1997; Sen 2004, and many others) are a result of misidentifications of specimens by authors who have not compared their material with specimens of / literature that provide a morphological description of M. montana s.s. (i.e., no NE Indian Megophrys species possess the large supraocular horn like projections and dorsal ridge configuration of M. montana), citing papers that have misidentified specimens, or incorrect interpretation of historical taxonomic literature discussing the species. Dubois (1982, 1989) demonstrated that M. monticola Kuhl and Van Hasselt was an incorrect subsequent spelling of M. montana Kuhl and Van Hasselt. Considering Xenophrys monticola and Megophrys parva as conspecifics, Dubois (1992) made a case to the International Commission of Zoological Nomenclature (ICZN) to formally suppress Xenophrys monticola Günther for the purposes of the Principle of Priority (but not for those of the Principle of Homonymy), to retain M. parva due to prevailing usage as the valid name and thus prevent perceived confusion regarding historical literature using the species name “ monticola ”. The ICZN took a formal action on this request in Opinion 1763 (Anonymous 1994), and added “ Xenophrys monticola Günther ” to the Official Index of Rejected and Invalid Specific Names in Zoology.</p><p>Though the type series of M. monticola and M. parva are superficially similar in general appearance, Mahony et al. (2017) indicated that M. parva s.s. is very likely to represent a member of the Southeast Asian endemic M. (Xenophrys) lekaguli species group (Figure 4; Appendix II, Figures 5 &amp; 6) based on the molecular systematic position of specimens collected relatively nearby the type locality of M. parva, and that were considered to be morphologically similar to the type specimens. Therefore M. parva s.s. is not conspecific with X. monticola s.s. With this information, it is now obvious that few historical reports of X. “ monticola ” or M. “ parva ” represent M. monticola s.s. as taxonomically redefined here. Moreover, the name M. montana Kuhl and Van Hasselt (/ M. monticola Kuhl &amp; Van Hasselt) has also been widely used in literature for several other species (e.g., M. ligayae, M. nasuta, M. stejnegeri) which were recognised as subspecies (after Inger 1954) until relatively recently (e.g., Iskandar 1998). At no point in time, that we are aware of, have both monticola Günther (from Northeast India-Nepal) and the misspelled “ monticola ” Kuhl and Van Hasselt (from Java, Indonesia) been included in the same genus, demonstrating that taxonomic rather than nomenclatural confusion is the primary issue in historical literature. So, we disagree with Dubois (1982, 1989) that widespread confusion from historical literature may be caused by the recognition of M. monticola (Günther) as a valid name. To this effect, an application to revive the combination Xenophrys monticola Günther to represent a distinct species-level taxon will be submitted to the ICZN.</p><p>Deuti et al. (2017) described two new species, Megophrys sanu comb. nov. (type locality: “Latpanchar [26°54'34"N, 88°23'53"E, 1110 m], Darjeeling district, West Bengal, India ”) and Megophrys katabhako comb. nov. (type locality: “Kabi [27°41'23"N, 88°62'82"E, 1410 m], North district, Sikkim, India ”). It should be noted that two of three GPS coordinates for tissue sampled specimens provided by Deuti et al. (2017) were found to be erroneous during the preparation of our map (Figure 8B), i.e., their longitude coordinates for Kabi were technically not possible since 60 minutes is the maximum in DMS (degrees, minutes, seconds) format: the correct coordinates for Kabi village is ~ 27°24'21"N, 88°37'03"E. Likewise, a stream near Senchal Lake (given as “ 27°06'02"N, 88°17'00"E ”) for ZSIC A 11799 lies nearby Jorethang, Sikkim state, almost 15 km north of the lake (Senchal Lake is at ~ 26°59'37"N, 88°15'54"E). The two new species were recognised based on a morphological examination of recent collections, a comparison with the type specimens of M. monticola and M. parva (amongst others), and DNA barcoding using a short fragment of 16S DNA. Besides those mentioned above, a number of additional errors/oversights in Deuti et al. (2017) are clearly evident, the most obvious of which require discussion here in order to clarify the taxonomic status of their new species. In their molecular analyses, Deuti et al. (2017) included sequences of most of the taxa from the Chen et al. (2017) study except M. zhangi from Tibet. Unusually, M. zhangi appears to also be the only conspecific omitted from the morphological comparisons sections of both Megophrys sanu comb. nov. and M. katabhako comb. nov. (Deuti et al. 2017). We performed a nucleotide BLAST search of the 16S sequences of Megophrys sanu comb. nov. generated by Deuti et al. (2017) (GenBank numbers KX894678 –80) and found that they were 99–100% identical to Chen et al. ’s (2017) M. zhangi sequences, and thus are the same as our “mid-elevation” populations associated with M. monticola in this study (refer to the Systematic position section above). We also performed a nucleotide BLAST search of our high-elevation population of M. monticola (GenBank number KY 022312 from Mahony et al. 2017) and found 98% similarity with the holotype of M. katabhako comb. nov. To further demonstrate the phylogenetic affinities of these two taxa we ran a RAxML analysis that included sequences from Deuti et al. (2017), and additional 12S/12S-tVal-16S sequences we generated for M. monticola populations from the Darjeeling area (Dataset K; Appendix I, Table 2). This analysis demonstrated that M. katabhako comb. nov. and M. sanu comb. nov. are nested within our concept of M. monticola (Appendix II, Figure 7).</p><p>Measurements of male specimens in this study that genetically corresponded with M. katabhako comb. nov. and M. sanu comb. nov. were not comparable to those provided by Deuti et al. (2017), e.g., the adult male SVL ranges for both species were larger: M. katabhako comb. nov.: 35.0– 37.4 mm N = 3 in Deuti et al. (2017 ––note: despite stating twice in text that four adult males were available, without explanation they provided data for only three in their table 2, Principle Component Analysis, &amp; maybe other statistical analyses) vs. 38.2–42.3 mm N = 7 in our study; M. sanu comb. nov.: 39.0– 46.7 mm N = 5 in Deuti et al. (2017) vs. 42.4–49.5 mm N = 9 in our study. If considered separately, the SVL measurements from our study and Deuti et al. ’s (2017) do correspond with their claim that the two haplotypes differ from each other by adult male body size. In contrast, we found our “high-elevation” haplotype (equivalent to M. katabhako comb. nov.) to have distinctly shorter relative tibia length (average TL/SVL 46.4%, N =7) compared to our “mid-elevation” population (equivalent to M. sanu comb. nov.) (average TL/SVL 49.5% N =9), which is the opposite case reported by Deuti et al. (2017). Keratinised spinules are absent on the ventral surfaces of thighs on all specimens of both “high-” and “mid-elevation” populations in our study, indicating that this character is not diagnostic for M. katabhako comb. nov. We also did not find their stated differences in dorsal colouration and markings to be diagnostic. Further morphometric comparison between the specimens studied in Deuti et al. (2017) and this study is not possible because the authors did not provide measurements for individual specimens, but provided only range, mean and standard deviation for their specimen series.</p><p>The morphological differences observed between the series of specimen examined in Deuti et al. (2017) and our study might be the result of sample bias, or differences in measurement techniques/character delineation. However, Deuti et al. (2017) included specimens of two “high-elevation” populations (Ghoom [2448 m] &amp; Kolakham [1860 m]) in their concept of M. sanu comb. nov. without providing supporting molecular data. Their molecular data and ours both associate populations from the vicinity of Ghoom with their M. katabhako comb. nov. so it is possible that their specimen series of M. sanu comb. nov. contains both taxa. Due to the ambiguity of morphologically associating specimens to one or the other of these two mitochondrial haplotypes, and based on our more extensive molecular sampling which suggests elevational segregation of the haplotypes, we recommend that the haplotype association of specimens from localities not sampled molecularly in Deuti et al. (2017) and elsewhere, should be considered unknown.</p><p>Deuti et al. (2017), presumably unaware of the lectotypification made by Mahony et al. (2017), lectotypified the “Khasya” syntype (BMNH [18]53.8.12.52) of M. monticola . Since the publication date of Deuti et al. (2017; 31 st July 2017) postdates the publication date of Mahony et al. (2017; 18 th January 2017), the valid lectotype for Xenophrys monticola is that designated by Mahony et al. (2017) (BMNH [18]53.8.12.52 from “ Sikkim ”), according to Article 74.1.1 of the International Code of Zoological Nomenclature, hereafter referred to as the “ Code ” (ICZN 1999). Without any morphological justification, they assigned the Sikkim syntype (now lectotype) of M. monticola to their referred specimens of M. sanu comb. nov. If their identification was correct, then Megophrys sanu comb. nov. would be considered to represent a junior subjective synonym of Megophrys monticola (Günther, 1864) .</p><p>Deuti et al. (2017) referred to the lectotype of X. monticola as a juvenile female; however, both large and small ova are clearly present in the ovaries, thus we regard this specimen to be an adult. This individual is smaller (SVL 40.5 mm) than adult females of both M. “ katabhako ” comb. nov. (SVL 47.3–48.7 mm, N =2, this study) and M. “ sanu ” comb. nov. (SVL 56.1 mm, N =1, this study; SVL 41.4–60.2 mm, N =5, Deuti et al. 2017:table 2 [note: they list only two adult females in their “Specimens examined” and “Specimens allocated to new species” sections so it is unclear what five ‘adult’ specimens were included in the table]). Based on the available material in our study, the character we found that usually diagnosed the mitochondrial haplotypes is that FIL is less than FIVL on nine of the 10 specimens from our “high-elevation” haplotype (equivalent to M. katabhako comb. nov.) and the M. monticola lectotype. Whereas, on all 10 specimens from our “mid-elevation” haplotype (equivalent to M. sanu comb. nov.) and one specimen of our “high-elevation” haplotype, we found FIVL is less than FIL. We therefore suggest that if the M. monticola lectotype is truly conspecific with either of the two new species described by Deuti et al. (2017), based on morphology alone we consider it to be more similar to the smaller species M. katabhako comb. nov., than to M. sanu comb. nov. We therefore consider Xenophrys katabhako Deuti et al. 2017 to represent a junior subjective synonym of Xenophrys monticola Günther, 1864 .</p><p>Our analyses of nuclear markers from one specimen of each of the two mitochondrial haplotypes referring to M. katabhako comb. nov. and M. sanu comb. nov., found no significant divergence between these taxa (see Systematic position section above). An increased mitochondrial and nuclear DNA sampling of both mitochondrial haplotypes from across their ranges will help identify whether our results are an artefact of mitochondrial introgression between two species, e.g., Chen et al. ’s (2017) M. “ zhangi ” and M. monticola s.s., and whether M. monticola as redefined here, represents a single biological species. Therefore, in the absence of evidence suggesting otherwise, and pending further investigations into the taxonomic affinities of populations of small sized Megophrys species that occur west of Bhutan, we recommend that Xenophrys sanu Deuti et al., 2017 and Xenophrys katabhako Deuti et al., 2017 should both be considered as junior subjective synonyms of Megophrys monticola (Günther, 1864) .</p></div>	https://treatment.plazi.org/id/03D6878AFFD1022FFF73F894FE16FE39	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
03D6878AFFC3022FFF73FDEFFB43FA3E.text	03D6878AFFC3022FFF73FDEFFB43FA3E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megophrys (Xenophrys) zhangi Ye and Fei 1992	<div><p>Megophrys (Xenophrys) zhangi Ye and Fei, 1992</p><p>(Figure 10)</p><p>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.</p><p>Holotype. Adult male (CIB 750296: Figure 10), from “Zhangmo, Nyanang, Xizang altitude 1000 m ” (= <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=85.98&amp;materialsCitation.latitude=27.99" title="Search Plazi for locations around (long 85.98/lat 27.99)">Zhangmu town</a>, ~ 27°59'24"N, 85°58'48"E, Nyalam County, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=85.98&amp;materialsCitation.latitude=27.99" title="Search Plazi for locations around (long 85.98/lat 27.99)">Shigatse Prefecture</a>, Tibet [or Xizang] Autonomous Region, China), collected by Yongzu Zhang on 26 June 1975 (Ye &amp; Fei 1992).</p><p>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) .</p><p>Examined specimen. Adult male (CIB 750296––photos only), holotype.</p><p>Description of type series. Refer to Ye and Fei (1992) for the description in Chinese language text.</p><p>Morphological comparison. Adult body size range provided for Megophrys zhangi is from the original description (Ye &amp; Fei 1992).</p><p>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.</p><p>Systematic position. Refer to the Systematic position section for Megophrys monticola above.</p><p>Etymology. The specific epithet “ zhangi ” is a patronym, named for Yongzu Zhang, the collector of the type series.</p><p>Suggested common name: Zhang’s Horned Frog.</p><p>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.</p></div>	https://treatment.plazi.org/id/03D6878AFFC3022FFF73FDEFFB43FA3E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
03D6878AFFC30216FF73F9E8FCFAFCD8.text	03D6878AFFC30216FF73F9E8FCFAFCD8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megophrys (Xenophrys) robusta Boulenger 1908	<div><p>Megophrys (Xenophrys) robusta Boulenger, 1908</p><p>(Figures 11 &amp; 12; Table 1)</p><p>[?] Xenophrys gigas Jerdon 1870:85 (partim: “ Sikim ”). In: Notes on Indian herpetology. Proceedings of the Asiatic Society of Bengal, March, 1870: 66–85.</p><p>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.</p><p>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.</p><p>Paralectotypes (by implication). Two adult females (ZSIC 9681, ZSIC 10777), details are the same as for the lectotype .</p><p>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 &amp; F), from near <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Latpanchar town</a> (26°54'34.2"N, 88°23'54.54"E, 1135 m asl.), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Kurseong</a> sub-division, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Darjeeling district</a>, West Bengal, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Northeast</a> India, collected by the <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Systematics Lab</a> 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 <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Stream</a> 1 (26°54'34.7"N, 88°23'52.7"E, 1030 m asl.), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Mahananda Wildlife Sanctuary</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Latpanchar township</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Kurseong</a> sub-division, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Darjeeling district</a>, West Bengal, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Northeast</a> 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 <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Sessa village</a> (27°6'4.02"N, 92°31'38.52"E, 1110 m asl.), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">West Kameng district</a>, Arunachal Pradesh state, Northeast India, collected by the Systematics Lab members on 07–09 August 2009 .</p><p>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.</p><p>Forelimbs moderately long, thin (Figure 11A &amp; 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&lt;I=II&lt;III (FIL 12.5, FIIL 12.5, FIIIL 18.2, FIVL 12.2); interdigital webbing and lateral fringes on digits absent; subarticular, supernumerary and metacarpal tubercles absent; thenar tubercle weakly developed; digit tips slightly expanded, flattened, without pads or terminal grooves. Hindlimbs relatively long, thin (Figure 11A &amp; B); thighs slightly shorter than shanks, and feet (TL 50.0, SHL 52.6, FOL 50.8); toes long, thin (Figure 11E), relative toe lengths I&lt;II&lt;V&lt;III&lt;IV; digit tips slightly dilated, flattened, without distinct pads; base of digits with thick rudimentary webbing; lateral fringes on toes absent; outer metatarsal, subarticular and supernumerary tubercles all absent; inner metatarsal tubercle present but barely distinguishable; ridge of thickened skin on ventral surface of digits absent.</p><p>Skin on dorsal surfaces of snout, head, and back weakly granular; throat, chest, and limbs smooth; flanks densely granular with ~15 small scattered tubercles (counted on left side); entire abdomen wrinkled (artefact of preservation), posterior thighs and cloacal region finely granular; tympanum smooth, slightly concave; sides of head finely granular; palpebral horn absent but with a broad fleshy bump in its place; supratympanic ridges narrow anteriorly, widening posteriorly, extends from posterior orbital border, curves down along upper and posterior border of tympanum, terminating above forelimb insertion; narrow, raised dorsolateral ridge on each side, extending posteriorly from behind supratympanic ridge to ~75% trunk length; weak, V-shaped parietoscapular ridge present, extending posteriorly from temporal region on each side, meeting medially beyond level of forelimbs, second short weak inverted U-shaped sacral ridge on posterior dorsum; pectoral glands weakly raised, level with axilla on chest; femoral glands not distinctly raised, on posterior surface of thighs, closer to knee than to cloaca; dermal asperities absent.</p><p>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).</p><p>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 HWŽHL (this study, N =9; Deuti &amp; Kumar 2008, N =5), Arunachal Pradesh specimens have HLŽHW (this study, N =3); two specimens (SDBDU 2009.1270, SDBDU 2009.1245) have relative finger length formula IV=I=II&lt;III; on most specimens, vomerine ridges are positioned equidistant from choanae and each other; vomerine teeth vary in size considerably, from indistinct on approximately half of examined specimens, to moderately well-developed on SDBDU 2011.1057; rounded appearance of tongue on lectotype appears to be artefact of preservation, as tongue of other specimens are distinctly, though weakly bifurcate posteriorly; dorsolateral ridges typically extend posteriorly almost to above cloaca on most specimens; tympanic concealment by supratympanic ridge varies from ~10 to 40%; parietoscapular-sacral ridge configurations vary considerably, i.e., “&gt; (”, “&gt;--&lt;”, or only V-shaped parietoscapular ridge present; dermal asperities absent on all females and subadults, but all males from Darjeeling possess black spinular asperities in dense thick circummarginal band along lower and upper jaw, covering entire surface below (and including) supratympanic ridges, more sparse on loreal region, absent from front of snout, sparse on posterior surface of upper eyelids, posterior dorsal surface of head and anterior body, increasing in density posteriorly on dorsum, sparse on dorsal and ventral surfaces of thighs, shanks, and tarsi, but more dense on dorsal surfaces near joints, sparse (or occasionally absent) on posterior surface of abdomen, present along all dorsal ridges, absent from flanks, anterior abdomen, chest and throat. Males from Arunachal Pradesh agree with the Darjeeling specimens regarding asperities cover, except that they have less dense asperities on upper jaw, none on supratympanic ridges except on tympanum and surface posterior to tympanum. Some specimens have weakly developed outer metacarpal tubercles visible; moderately well developed thenar tubercles present on some individuals. Flank tuberculation varies from sparse, small tubercles to large, dense tubercles. Dorsal surfaces of snout usually with small irregularly arranged dark brown blotches (Figure 12A, B &amp; D); triangular marking on dorsal surface of head occasionally incomplete; parietoscapular-sacral ridges typically accompanied by dark brown marking that follows shape of dermal ridges, others densely mottled with brown blotches over entire dorsum; general colouration in preservation ranges from pale brown to dark brown dorsally, dorsal marking barely visible (or absent) on darker individuals; some specimens have 3–6 distinct light brown longitudinal stripes extending from gular region onto chest (Figure 12E &amp; F); most have faint to distinct brown blotch extending from posterior border of orbit through tympanum (Figure 12A &amp; B), and from anterior orbit laterally along edge of canthus rostralis to snout tip; all have distinctly dark brown lower edge to supratympanic ridge and brown vertical bars on sides of face, most distinct is broad bar extending from lower border of orbits to edge of upper lip [erroneously reported to be absent in M. robusta by Ohler et al. (2002)] (Figures 11C, &amp; 12A, C &amp; E); outer edge of gular region typically brown, with eight light patches.</p><p>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.</p><p>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.</p><p>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 &amp; 3; Appendix II, Figures 2, 3A &amp; 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 &amp; 6). Refer to Appendix I, Table 6 for uncorrected p -distances for the 16S rRNA gene between M. robusta and other MMSG species.</p><p>Etymology. The specific epithet “ robusta ” is Latin for “hard” or “strong”, presumably a reference to the heavy set habitus of this species.</p><p>Suggested common name: Robust Horned Frog.</p><p>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.</p><p>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 &amp; 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.</p><p>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 &amp; 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.</p><p>Megophrys robusta has been reported from several localities further east in Arunachal Pradesh state, e.g., Lower Subansiri district (Bordoloi et al. 2000; Bordoloi &amp; Borah 2001) and the Upper Siang district and Changlang districts (Pawar &amp; 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 .</p><p>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 species present in Meghalaya. An early report of M. robusta from the East Khasi Hills district, Meghalaya by Hora (1923) was based solely on nondescript tadpoles, and thus this locality should not be included in the species distribution.</p><p>Khan’s (2008) inclusion of M. robusta on a checklist for Bangladesh (from “NE and N of hills”) herpetofauna is likely erroneous based on geographical distance from verified populations. Furthermore, within the political borders of Bangladesh the maximum elevation barely exceeds 1000 m asl., less than the lowest recorded elevation verified for this species. For the distribution of M. robusta, Sen and Mathew (2008) listed, aside from Arunachal Pradesh state, the Northeast Indian states of Assam, Meghalaya and Nagaland, and the countries of Bangladesh, China (and specifically Hong Kong), Myanmar and Vietnam without accompanying justification, hence these uncorroborated localities are at best considered erroneous.</p></div>	https://treatment.plazi.org/id/03D6878AFFC30216FF73F9E8FCFAFCD8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
03D6878AFFFA0214FF73FCB5FE09FD48.text	03D6878AFFFA0214FF73FCB5FE09FD48.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megophrys (Xenophrys) medogensis Fei, Ye and Huang 1983	<div><p>Megophrys (Xenophrys) medogensis Fei, Ye and Huang, 1983</p><p>(Figure 13)</p><p>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.</p><p>Holotype. Adult male (CIB 73 II0015: Figure 13), from “Motuo, Xizang, alt. 1000m ” (= <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=95.340836&amp;materialsCitation.latitude=29.332224" title="Search Plazi for locations around (long 95.340836/lat 29.332224)">Medog</a> [or <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=95.340836&amp;materialsCitation.latitude=29.332224" title="Search Plazi for locations around (long 95.340836/lat 29.332224)">Motuo</a>] town [~ 29°19'56"N, 95°20'27"E], Medog County, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=95.340836&amp;materialsCitation.latitude=29.332224" title="Search Plazi for locations around (long 95.340836/lat 29.332224)">Nyingtri Prefecture</a>, Tibet [or Xizang] <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=95.340836&amp;materialsCitation.latitude=29.332224" title="Search Plazi for locations around (long 95.340836/lat 29.332224)">Autonomous Region</a>, China), collectors presumably <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=95.340836&amp;materialsCitation.latitude=29.332224" title="Search Plazi for locations around (long 95.340836/lat 29.332224)">Fei</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=95.340836&amp;materialsCitation.latitude=29.332224" title="Search Plazi for locations around (long 95.340836/lat 29.332224)">Hu</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=95.340836&amp;materialsCitation.latitude=29.332224" title="Search Plazi for locations around (long 95.340836/lat 29.332224)">Ye</a> and Wu on 17 July 1973 (Fei et al. 1977).</p><p>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) .</p><p>Examined specimens. Adult male (CIB 73II 0015: images only: Figure 13), holotype.</p><p>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.</p><p>Systematic position. Refer to the Systematic position section above for M. robusta .</p><p>Morphological comparison. Adult body size ranges provided for M. medogensis are from Fei et al. (2009) (adult males, N =16).</p><p>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.</p><p>Etymology. The specific epithet “ medogensis ” is a toponym, derived from the type locality of Medog county.</p><p>Suggested common name: Medog Horned Frog.</p><p>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 &amp; 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.</p><p>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.</p><p>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 &amp; 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.</p><p>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).</p></div>	https://treatment.plazi.org/id/03D6878AFFFA0214FF73FCB5FE09FD48	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
03D6878AFFF80214FF73FD05FAE7FC78.text	03D6878AFFF80214FF73FD05FAE7FC78.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megophrys (Xenophrys) major Mahony & Kamei & Teeling 2018	<div><p>Megophrys (Xenophrys) major s.s. Boulenger, 1908</p><p>(Figures 14 &amp; 15; Table 1)</p><p>Xenophrys gigas Jerdon 1870:85 (partim: Khasi Hills). In: Notes on Indian herpetology. Proceedings of the Asiatic Society of Bengal, March, 1870: 66–85.</p><p>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.</p></div>	https://treatment.plazi.org/id/03D6878AFFF80214FF73FD05FAE7FC78	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
03D6878AFFF8021FFF73FC33FBF2FEA8.text	03D6878AFFF8021FFF73FC33FBF2FEA8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Xenophrys gigas Jerdon 1870	<div><p>Neotype for Xenophrys gigas Jerdon, 1870 and Megalophrys major Boulenger, 1908 (by present designation).</p><p>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 &amp; B), from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=93.642494&amp;materialsCitation.latitude=24.86111" title="Search Plazi for locations around (long 93.642494/lat 24.86111)">Raenghkucpaeng</a> duithuak (duithuak = stream, in local <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=93.642494&amp;materialsCitation.latitude=24.86111" title="Search Plazi for locations around (long 93.642494/lat 24.86111)">Rongmei</a> [<a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=93.642494&amp;materialsCitation.latitude=24.86111" title="Search Plazi for locations around (long 93.642494/lat 24.86111)">Nruangmei</a>] language), near <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=93.642494&amp;materialsCitation.latitude=24.86111" title="Search Plazi for locations around (long 93.642494/lat 24.86111)">St. Joseph School</a> (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 &amp; D), from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=93.623055&amp;materialsCitation.latitude=24.85" title="Search Plazi for locations around (long 93.623055/lat 24.85)">Mpeih</a> 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 &amp; F), from near <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=93.498055&amp;materialsCitation.latitude=24.966946" title="Search Plazi for locations around (long 93.498055/lat 24.966946)">Khangchiuluanh</a> (<a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=93.498055&amp;materialsCitation.latitude=24.966946" title="Search Plazi for locations around (long 93.498055/lat 24.966946)">Khongjaron</a>) <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=93.498055&amp;materialsCitation.latitude=24.966946" title="Search Plazi for locations around (long 93.498055/lat 24.966946)">Church</a> (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 .</p><p>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.</p><p>Forelimbs long, thin (Figure 14A &amp; 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&lt;II=I&lt;III (FIL 9.3, FIIL 9.3, FIIIL 12.0, FIVL 9.1); interdigital webbing, subarticular, supernumerary and metacarpal tubercles all absent; thenar tubercles weakly developed; finger tips flattened, slightly expanded relative to digit widths (FIIIW 2.0, FIIIDW 2.3) with subcircular pads, terminal grooves on pads absent. Hindlimbs long, thin (Figure 14A &amp; B); thighs slightly shorter than shanks, and longer than feet (TL 39.7, SHL 40.8, FOL 37.1); toes long, dorsoventrally flattened with very narrow lateral fringes (Figure 14E), relative toe lengths I&lt;II&lt;V&lt;III&lt;IV; toe tips flattened, considerably dilated (TIVW 2.0, TIVDW 2.7), with distinct subcircular pads, terminal grooves on pads absent; webbing basal, I1.5– 2II 1–2.5 III2 –3.5IV3.5– 2V; inner metatarsal tubercles distinct (IMT 4.5), longitudinally ovoid; subarticular, supernumerary and outer metatarsal tubercles absent; ridge of callous tissue absent on ventral surface of toes.</p><p>Skin of dorsal surfaces of head and body primarily smooth with weak, densely scattered granules; tympanum with borders raised relative to surrounding region; outer edge of upper eyelid with a distinct pointed bump; supratympanic ridges narrow anteriorly, gradually expanding posterior to tympanum to become moderately enlarged and glandular, extending from posterior orbital borders, broadly curving through upper border of tympanum, terminating above forearm insertions; flanks densely covered with small to large unevenly scattered pustular tubercles; dorsolateral ridges thin, well defined, extending from behind supratympanic ridge to ~70% distance to vent; parietoscapular-sacral ridges weak, “&gt;–&lt;” configuration, composed primarily of single row of closely spaced asperities; dorsal surfaces of forearms, thighs and shanks with short transverse ridges; posterior thighs with small scattered pustular tubercles, remaining dorsal surfaces of forelimbs and hindlimbs otherwise smooth; gular region, chest, abdomen and ventral surfaces of limbs smooth; pectoral glands small, raised, level with axilla on chest; femoral glands large, flat, on posterior surface of thighs, slightly closer to knee than to cloaca; distinct pair of enlarged glands present adjacent to cloaca, one on each side, that are smaller than femoral glands but larger than surrounding tubercles; small black and white dermal asperities present, forming broad dense band circummarginally on gular region, dense on upper lips, lower loreal region, and tympanic region (especially dense on tympanum), moderately dense on dorsal surface of eyelids, entire dorsal surface of head and back, increasing in density posteriorly and on dorsal ridges, absent from all remaining surfaces.</p><p>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.</p><p>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, IV&lt;II=I&lt;III on all “Darjeeling” specimens except a subadult (BMNH 1947.2.24.99) with IV=II=I&lt;III, all Sohra specimens have IV=II=I&lt;III, except a female (ZSIC 9746) with IV&lt;II&lt;I&lt;III, and all aforementioned variations, with an additional IV&lt;I&lt;II&lt;III for RGK 0 0 98, are observed in the Nagaland-Manipur specimens; vomerine ridges vary from sub-circular to ovoid, positioned slightly closer to choanae than to each other on many individuals; posterior edge of tongue on some specimens appears more deeply notched than others, likely due to preservation condition; dorsolateral ridges typically weak to moderately well developed, varying in length from ~60 to 90% trunk length; between ~5 and 30% of tympanum can be concealed by supratympanic ridge; dermal tubercle cover on flanks varies considerably, some with only sparse scattering of small tubercles (e.g., Figure 15A), others with moderately dense cover of heterogeneous sized (large to small) tubercles (e.g., Figure 15C &amp; E); dorsal markings vary from faint to distinct complete X-shaped, Yshaped, hourglass-shaped marking, or just a V-shaped marking on parietoscapular region, not visible on some older specimens presumably due to fading; some specimens have faint spotting and blotching on chest and abdomen (e.g., Figure 15F); density and surface coverage of dermal asperities varies extensively among male specimens, if present on females, asperities sparse on mid-dorsum increasing in density posteriorly to above cloaca, but absent from other surfaces, asperities typically absent on juvenile and subadult specimens examined; white upper lip stripe not present on metamorphs and juveniles (with SVL &lt;50 mm), instead lateral surfaces of head plain dark brown.</p><p>Secondary 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.</p><p>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.</p><p>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 &amp; 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).</p><p>Etymology. The original species epithet “ gigas ” is derived from the Greek word meaning “giant”, later replaced with “ major ”, Latin for “large”.</p><p>Suggested common name: Jerdon’s White-lipped Horned Frog.</p><p>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.</p><p>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 &amp; 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).</p><p>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.</p><p>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 “large”, presumably relative to the considerably smaller Xenophrys monticola, the only other Xenophrys species known at that time; 3) ZSIC 10777 and ZSIC 9681 are M. robusta (SM per. obs.); 4) Sclater (1892a, 1892b) provided a list of types in the Indian Museum that included only one megophryid, Scutiger (as Cophophryne) sikkimensis (Blyth, 1855), and Sclater (1892a) explicitly stated that the collections then contained the types of only one of Jerdon’s species, Hyla annectans (Jerdon, 1870) .</p><p>Before leaving India, Jerdon visited and collected specimens in and around Darjeeling–Sikkim and the Khasi Hills, but he notoriously did not maintain notes on his collections, relying largely on memory for details of localities (Günther 1875). Upon his return to England, Jerdon assisted Günther with the accessioning of his collection into the BMNH– –“The work of systematically arranging and naming this collection was carried on jointly by him [Jerdon] and myself, and proceeded as far as the genus Tropidonotus, when it was interrupted by an illness from which he never recovered….and I am therefore ignorant of the habitat [collection localities] of a part of the specimens which were still unexamined at the time of his death” (Günther 1875). Jerdon’s original “ X. gigas ” collection consisted of a series of presumably 17 specimens (BMNH [18]72.4.17.398–414), and was one of the last taxa from his collection to be accessioned (BMNH Accession Register). The BMNH Accession Register gives the locality “Darjeeling (&amp; Khasia)”, but the locality appeared in subsequent literature as Darjeeling only (e.g., Boulenger 1882 [“h”]). Doubts regarding whether some of Jerdon’s other herpetological specimens were from the Khasi Hills or from Darjeeling have been noted elsewhere (e.g., the type locality of Ixalus jerdoni Günther, 1876 [= Frankixalus jerdoni] from “Darjeeling” was subsequently found to be widespread in the Khasi Hills [Biju et al. 2016] but not reported from Darjeeling or surrounding regions since its original description). Jerdon deposited most (if not all) of his type specimens in the British Museum and Indian Museum, but since the Indian Museum did not (according to Sclater [1891a, 1891b]) contain Jerdon’s megophryid type specimens, the BMNH specimen series may have contained some, or all, of the five original specimens based on which Jerdon (1870) coined the name.</p><p>Remarks on Megophrys major Boulenger, 1908: Xenophrys gigas was subsequently transferred to the genus Megalophrys by Boulenger (1908) rendering Megalophrys gigas (Jerdon, 1870) a junior homonym of Megalophrys gigas Blyth, 1855 [now Nanorana liebigii (Günther, 1860)]. To correct this, Boulenger (1908) coined the replacement name Megalophrys major, and provided a morphological description of the species with a note that Jerdon (1870) named X. gigas based on specimens from Darjeeling and the Khasi Hills “now preserved in the British Museum [now NHMUK] and Indian Museum [now ZSIC]”. Considering Sclater (1892a, 1892b) did not recognise types for this species in the Indian Museum, and up until Boulenger (1908), none of Jerdon’s NHMUK Megophrys specimens were recognised as “types” of X. gigas (e.g., Günther 1876; Boulenger 1882:442), the provenance of Boulenger’s statement is unclear. In Boulenger’s (1908) account of X. gigas (as M. major), he identified two specimens (an adult male and an adult female– –accession numbers not given) as “type” for which he provided measurements, but parts of his morphological description clearly indicated that he included many other specimens. It is unclear whether Boulenger regarded the entire specimen series to be types, or only the two specimens mentioned, however, in the absence of stringent and universally accepted nomenclatural rules governing typification at that time, Boulenger may have designated specimens collected by Jerdon as types as an attempt to remove further ambiguity. The BMNH Specimen Catalogue currently lists all 17 of Jerdon’s original specimen series as “ syntypes ” of M. major, but there is no indication in the BMNH Accessions Register that these specimens were originally considered to be the types of X. gigas . According to the Code (Article 72.7), a replacement name is an objective synonym, and must have the same type series as the original name, therefore, in the absence of evidence that Boulenger’s “type[s]” represent two of the five syntypes of X. gigas, his typification should be considered invalid.</p><p>Currently, half of the NHMUK “ syntype ” series (BMNH [18]72.4.17.407–414) are no longer present in the collection and must be assumed to have been exchanged with other museum collections prior to the 1947 replacement numbers (e.g., MCZ A-6814, received on exchange from the “British Museum” in 1920 [MCZbase, online catalogue, accessed 20 January 2018]). The remaining adult specimens in the “ syntype ” series in the NHMUK appear to be homogeneous, however, the specific status of the subadult female and juvenile specimens cannot be confirmed based on morphological characters alone. Aside from Jerdon’s NHMUK specimen series, all of which were regarded by Günther (1872) to have been collected from Darjeeling, we are aware of no subsequent collections of M. major s.s. from north of the Brahmaputra river– – Daniel (1962) reported M. major from Darjeeling based on two specimens but the figured animal clearly portrayed a M. robusta, and Subba et al. 2016 did not observe this species in Sikkim. However, we demonstrate that M. major s.s. is present in the southern Khasi Hills and adjacent areas, thus it more likely that Jerdon’s NHMUK specimen series came from there. Until the time specimens referable to M. major s.s. are collected from Darjeeling/ Sikkim area (which we consider unlikely), we regard Jerdon’s NHMUK specimen series to have been collected from the Khasi Hills, Meghalaya state, Northeast India. Likewise, we confirm that M. robusta is only found west of the Brahmaputra (including the Darjeeling area). Therefore, the name X. gigas was coined based on more than one species ( M. robusta and one or more of the Khasi Hills-endemic MMC species). Since Jerdon deposited no specimens of M. robusta in the NHMUK, it is certain that the NHMUK “ syntype ” series cannot contain all of the original type specimens of X. gigas .</p><p>Proposed Nomenclatural Resolutions: Although the syntype specimens of Xenophrys gigas might not actually be lost or destroyed, the knowledge as to which (if any) of Jerdon’s specimens represent this series is effectively “lost” so that they are no longer unambiguously identifiable. Jerdon’s (1870) original description of Xenophrys gigas provided no diagnostic characters that can permit either the identification of an existing specimen as one of the syntypes, or the application of the name to a single biological species, so Xenophrys gigas is therefore considered a nomen dubium. To resolve the nomenclatural conundrum and bring nomenclatural stability to the species name by restricting its use to a single biological species, we propose that one of the NHMUK specimens (BMNH 1947.2.24.93 [rr. BMNH [18]72.4.17.399]––described above) collected by Jerdon is designated as the neotype of Xenophrys gigas . This action will reinstate the recognition of Xenophrys gigas Jerdon, 1870 as a primary objective synonym of Megalophrys major Boulenger, 1908, and will also preserve the nomen M. robusta for the large species of Megophrys from Darjeeling. The neotypification of this specimen will also restrict the type locality of X. gigas, and by implication that of M. major, to “Darjeeling”, corrected herein to “Khasi Hills” (which currently encompasses the East Khasi Hills and West Khasi Hills districts), Meghalaya state, Northeast India. By this action, the remaining NHMUK specimens in the series (BMNH 1947.2.24.92, BMNH 1947.2.24.94–99, BMNH 1947.2.25.1) and those in the ZSIC regarded by Chanda et al. (2000) to represent the syntype series of X. gigas will no longer be referred to as name-bearing type specimens for this taxon.</p></div>	https://treatment.plazi.org/id/03D6878AFFF8021FFF73FC33FBF2FEA8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
03D6878AFFF30203FF73FE65FE55FAB4.text	03D6878AFFF30203FF73FE65FE55FAB4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megophrys (Xenophrys) Mahony & Kamei & Teeling 2018	<div><p>Megophrys (Xenophrys) flavipunctata sp. nov.</p><p>(Figures 16 &amp; 17; Table 1)</p><p>Holotype. Adult male (BNHS 6040 [field no. SDBDU 2009.304]: Figures 16 &amp; 17B), from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=91.74611&amp;materialsCitation.latitude=25.443611" title="Search Plazi for locations around (long 91.74611/lat 25.443611)">Mawphlang Sacred Grove</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=91.74611&amp;materialsCitation.latitude=25.443611" title="Search Plazi for locations around (long 91.74611/lat 25.443611)">Mawphlang</a> (25°26'37"N, 91°44'46"E, 1810 m asl.), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=91.74611&amp;materialsCitation.latitude=25.443611" title="Search Plazi for locations around (long 91.74611/lat 25.443611)">East Khasi Hills district</a>, Meghalaya state, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=91.74611&amp;materialsCitation.latitude=25.443611" title="Search Plazi for locations around (long 91.74611/lat 25.443611)">Northeast</a> India, collected by members of the Systematics Lab, University of Delhi on 0 1 June 2009.</p><p>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 &amp; 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 .</p><p>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 &amp; F), from near <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=94.05862&amp;materialsCitation.latitude=25.639723" title="Search Plazi for locations around (long 94.05862/lat 25.639723)">Sazolie College</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=94.05862&amp;materialsCitation.latitude=25.639723" title="Search Plazi for locations around (long 94.05862/lat 25.639723)">Jotsoma village</a> (25°38'23"N, 94°03'31"E, 1575 m asl.), Kohima district, Nagaland state, Northeast India, collected by RGK on 27 June 2007 .</p><p>Holotype description (measurements in mm). Mature male (SVL 68.4) (Figures 16 &amp; 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.</p><p>Forelimbs moderately long, thin (Figure 16A &amp; 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&lt;I=II&lt;III (FIL 8.7, FIIL 8.6, FIIIL 11.9, FIVL 7.8); interdigital webbing, subarticular, supernumerary and metacarpal tubercles all absent; thenar tubercles weak; finger tips rounded, slightly expanded relative to digit widths (FIIIW 1.3, FIIIDW 1.6), longitudinally oval-shaped pads present, terminal grooves on pads absent. Hindlimbs long, thin (Figure 16A &amp; B); thighs slightly shorter than shanks, and slightly longer than feet (TL 35.5, SHL 36.7, FOL 34.7); toes long, dorsoventrally flattened, with narrow lateral fringes (Figure 16E), relative toe lengths I&lt;II&lt;V&lt;III&lt;IV; toe tips rounded, not dilated (TIVW 1.6, TIVDW 1.7), longitudinally ovalshaped pads distinct, terminal grooves on pads absent; webbing basal (III2.3–3.8IV3.5– 2V); inner metatarsal tubercle indistinct; subarticular, supernumerary and outer metatarsal tubercles absent; callous tissue weak on ventral surface of toes.</p><p>Skin of dorsal surfaces of body, limbs, and dorsal and lateral surfaces of head primarily smooth; tympanum smooth with borders slightly raised; outer edge of upper eyelid with a weak medial bump, remaining surface covered with small granular tubercles; supratympanic ridges narrow anteriorly gradually expanding beyond posterior edge of tympanum, becoming moderately enlarged, glandular, extending from posterior orbital border and broadly curving through upper border of tympanum, terminating above axilla; flanks with small to large scattered pustular tubercles; dorsolateral ridges thin, well defined, extending posteriorly from behind supratympanic ridges to ~75% trunk length; parietoscapular-sacral ridges indistinct; gular region, chest, abdomen and ventral surfaces of limbs smooth; pectoral glands faint, flat, level with axilla on chest; femoral glands large, flat, on posterior surface of thighs, slightly closer to knees than to cloaca; dermal asperities absent from all surfaces.</p><p>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.</p><p>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 II&lt;IV&lt;I&lt;III, and BNHS 6041 has the formula IV&lt;II&lt;I&lt;III; vomerine teeth indistinct on BNHS 6 043; all paratypes and referred specimens bar BNHS 6044 differ from holotype by possessing dorsolateral ridges that extend almost full length of trunk (e.g., Figure 17A, C &amp; E); supratympanic ridges conceal up to ~20% of upper tympanum; BNHS 6044 has very small black dermal asperities forming circummarginal band on gular region, few also present on flank tubercles, but absent from other surfaces; pectoral glands on some individuals small, raised; dorsal ridges vary from a weakly raised Vshaped parietoscapular ridge only (SDBDU 2009.297, SDBDU 2009.298, BNHS 6044, BNHS 6042), Y-shaped parietoscapular ridge either without (BNHS 6043), or with an attached inverted V-shaped sacral ridge on middorsum (all remaining specimens); BNHS 6041 and BNHS 6043 have moderately well developed thenar tubercles; BNHS 6043 in life was orange-brown dorsally with dense orange spotting through upper lip stripe and throat, spotting decreased in density posteriorly on abdomen (Figure 17C &amp; D); SDBDU 2009.297 in life had upper half of posterior supratympanic ridge creamish-white, blotches on central portion of throat and chest were a pale brownish-mauve; BNHS 6042 in life had chest, abdomen (excluding a large brown blotch and small speckling) and ventral surface of thighs creamish-white with distinct hint of yellow laterally on lower-most flanks; most specimens had distinct X-shaped marking on mid-dorsum.</p><p>Secondary 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.</p><p>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.</p><p>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).</p><p>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 &amp; 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.</p><p>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.</p><p>Suggested common name: Yellow Spotted White-lipped Horned Frog.</p><p>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.</p><p>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 &amp; 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 &amp; 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.</p><p>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.</p></div>	https://treatment.plazi.org/id/03D6878AFFF30203FF73FE65FE55FAB4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
03D6878AFFEF0207FF73FA1EFD52FAFC.text	03D6878AFFEF0207FF73FA1EFD52FAFC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megophrys (Xenophrys) Mahony & Kamei & Teeling 2018	<div><p>Megophrys (Xenophrys) oreocrypta sp. nov.</p><p>(Figures 18 &amp; 19; Table 1)</p><p>Holotype. One adult female (BNHS 6045 [field no. SDBDU 2009.1108]: Figures 18, &amp; 19A &amp; C), from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=90.22912&amp;materialsCitation.latitude=25.508167" title="Search Plazi for locations around (long 90.22912/lat 25.508167)">Tura Peak Reserve Forest</a> (25°30'29.4"N, 90°13'44.82"E, 640 m asl.), near <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=90.22912&amp;materialsCitation.latitude=25.508167" title="Search Plazi for locations around (long 90.22912/lat 25.508167)">Tura town</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=90.22912&amp;materialsCitation.latitude=25.508167" title="Search Plazi for locations around (long 90.22912/lat 25.508167)">West Garo Hills district</a>, Meghalaya state, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=90.22912&amp;materialsCitation.latitude=25.508167" title="Search Plazi for locations around (long 90.22912/lat 25.508167)">Northeast</a> India, collected by members of the Systematics Lab, University of Delhi on 20 June 2009.</p><p>Paratypes. Four unsexed juveniles (BNHS 6046 – BNHS 6049 [field nos. SDBDU 2009.1104 – 1107]: Figure 19B, D, E &amp; F), collected along with the holotype .</p><p>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.</p><p>Holotype description (measurements in mm). Mature female (SVL 94.9) (Figures 18, &amp; 19A &amp; 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.</p><p>Forelimbs moderately long, thin (Figures 18A &amp; B, &amp; 19A &amp; 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&lt;II&lt;I&lt;III (FIL 12.4, FIIL 10.7, FIIIL 15.4, FIVL 10.1); interdigital webbing, subarticular, supernumerary and metacarpal tubercles absent; thenar tubercles weak; finger tips rounded, slightly expanded relative to digit widths (FIIIW 1.8, FIIIDW 2.0), with subcircular pads, terminal grooves on pads absent. Hindlimbs long, thin (Figures 18A &amp; B, &amp; 19A &amp; C); thighs shorter than shanks, longer than feet (TL 44.8, SHL 48.0, FOL 41.6); toes long, dorsoventrally flattened, without lateral fringes (Figure 18E), relative toe lengths I&lt;II&lt;V&lt;III&lt;IV; toe tips rounded, moderately expanded relative to digit width (TIVW 2.0, TIVDW 2.5) with distinct longitudinally oval-shaped pads, terminal grooves on pads absent; webbing basal, to distal edge of metatarsal on Toe IV (3III 2.5– 4IV 4.2–2.5V); inner metatarsal tubercle very weakly developed; subarticular, supernumerary and outer metatarsal tubercles absent; ridge of callous tissue absent on ventral surface of toes.</p><p>Skin of dorsal surfaces of body, limbs, and dorsal and lateral surfaces of head primarily smooth; tympanum smooth, slightly concave, with borders slightly raised; outer edge of upper eyelids with a weak medial bump; supratympanic ridges narrow anteriorly gradually expanding beyond posterior edge of tympanum to become moderately enlarged and glandular, extending from posterior edge of orbits, through upper portion of tympanum, curving down abruptly at posterior tympanum border, terminating above forelimb insertion; flanks weakly granular with medium to large scattered pustular tubercles; dorsolateral ridges thin, weakly defined, extending from behind supratympanic ridges, almost entire length of trunk; weakly developed V-shaped parietoscapular ridge present, sacral ridges absent; ventral surfaces of head, body and limbs smooth; pectoral glands small, weakly raised, positioned on chest at mid-level of forelimb insertion; femoral glands large, flat, on posterior surface of thighs, slightly closer to knees than to cloaca; dermal asperities absent from all surfaces.</p><p>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 &amp; 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.</p><p>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 IV&lt;I=II&lt;III; posterior edge of tongue (examined on FMNH 74154 and BNHS 6046) distinctly bifurcate posteriorly; up to ~20% of upper portion of tympanum appears to be concealed by supratympanic ridge; dorsal surfaces of head and body of juveniles weakly granular; light upper lip stripe not visible on specimens smaller than SVL 41.0 mm (e.g., Figure 19E &amp; F); BNHS 6046 has distinct brown X-shaped marking on dorsum; BNHS 6046 and BNHS 6047 have small white asperities along dorsal ridges (e.g., Figure 19E). See Figure 19C and 19D for ventral colouration variation between holotype and juvenile specimens.</p><p>Secondary sexual characters. Males currently unknown.</p><p>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).</p><p>Systematic position. This taxon represents M. cf. major 1 in the molecular analyses (discussed as OTU 7), and in Mahony et al. (2017).</p><p>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 &amp; 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.</p><p>Etymology. The specific epithet “ oreocrypta ” is an adjective derived from the Latinised Greek words óros 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.</p><p>Suggested common name: Garo White-lipped Horned Frog. Sangma and Saikia (2015) provide the local Garo (language) name “Diplok Nakma” for this species.</p><p>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.</p><p>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.</p><p>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.</p></div>	https://treatment.plazi.org/id/03D6878AFFEF0207FF73FA1EFD52FAFC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
03D6878AFFEB0208FF73FA56FA8FFF3B.text	03D6878AFFEB0208FF73FA56FA8FFF3B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megophrys (Xenophrys) Mahony & Kamei & Teeling 2018	<div><p>Megophrys (Xenophrys) himalayana sp. nov.</p><p>(Figures 20 &amp; 21; Table 1)</p><p>Holotype. Adult male (BNHS 6050 [field no. SDBDU 2009.1227]: Figures 20, &amp; 21A, F &amp; H), from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.58061&amp;materialsCitation.latitude=27.082367" title="Search Plazi for locations around (long 92.58061/lat 27.082367)">Elephant village</a> (27°4'56.52"N, 92°34'50.22"E, 370 m asl.), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.58061&amp;materialsCitation.latitude=27.082367" title="Search Plazi for locations around (long 92.58061/lat 27.082367)">West Kameng district</a>, Arunachal Pradesh state, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.58061&amp;materialsCitation.latitude=27.082367" title="Search Plazi for locations around (long 92.58061/lat 27.082367)">Northeast</a> India, collected by members of the Systematics Lab, University of Delhi on either 0 4 or 0 5 August 2009.</p><p>Paratypes. Four adult males (BNHS 6051–6054 [field nos. SDBDU 2009.1206 – 1209]: Figure 21C, D, F &amp; 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 .</p><p>Referred specimens. Adult male (SDBDU 2009.787: Figure 21E &amp; G), and adult female (SDBDU 2009.750: Figure 21B &amp; 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.</p><p>Holotype description (measurements in mm). Mature male (SVL 72.1) (Figures 20, &amp; 21A, F &amp; 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.</p><p>Forelimbs long, thin (Figure 20A &amp; 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&lt;II&lt;I&lt;III (FIL 9.1, FIIL 8.7, FIIIL 12.2, FIVL 7.5); interdigital webbing, subarticular and supernumerary tubercles absent; thenar and outer metacarpal tubercles weakly developed; finger tips flattened, not expanded relative to digit widths, subcircular pads present, terminal grooves on pads absent. Hindlimbs long, thin (Figure 20A &amp; B); thighs slightly shorter than shanks, and longer than feet (TL 36.7, SHL 37.5, FOL 33.1); toes long, rounded, not dorsoventrally flattened, without lateral fringes (Figure 20E), relative toe lengths I&lt;II&lt;V&lt;III&lt;IV; toe tips flattened, not dilated but with distinct oval-shaped pads, terminal grooves on pads absent; webbing basal; inner metatarsal tubercle weakly defined, longitudinally oval-shaped; subarticular, supernumerary and outer metatarsal tubercles absent; ridge of callous tissue absent on ventral surface of toes.</p><p>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.</p><p>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 &amp; 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.</p><p>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&gt;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., “&gt;––&lt;”, “&gt; &lt;”, “&gt;– &lt;”, or only V-shaped parietoscapular ridge present; coverage of dermal asperities varies mostly in density between individual males from Elephant Village, in comparison with holotype, however, male from Rengging Village (SDBDU 2009.787) has much less coverage of asperities, having white asperities mostly restricted to posterior dorsum and dorsal ridges; Rengging Village female (SDBDU 2009.750) has few white asperities on posterior dorsum and on dorsal surfaces of thighs; outer metacarpal tubercles not visible on some specimens; tubercle cover on flanks varies considerably, some with only sparse scattering of small tubercles, others with moderately dense cover of heterogeneous sized (large to small) tubercles (Figure 21 A–E); dorsal and ventral markings vary considerably between individuals (see Figure 21 which represent extremes in variation).</p><p>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.</p><p>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.</p><p>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).</p><p>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 &amp; 4; Appendix I, Table 3; Appendix II, Figures 1, 2 &amp; 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.</p><p>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.</p><p>Suggested common name: Himalayan Horned Frog.</p><p>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.</p><p>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.</p></div>	https://treatment.plazi.org/id/03D6878AFFEB0208FF73FA56FA8FFF3B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
03D6878AFFE4020DFF73FE9EFD51FF19.text	03D6878AFFE4020DFF73FE9EFD51FF19.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megophrys (Xenophrys) Mahony & Kamei & Teeling 2018	<div><p>Megophrys (Xenophrys) periosa sp. nov.</p><p>(Figures 22 &amp; 23; Table 1)</p><p>Holotype. Adult male (BNHS 6055 [field no. SDBDU 2009.793]: Figures 22, &amp; 23A &amp; H), from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=94.986115&amp;materialsCitation.latitude=28.209435" title="Search Plazi for locations around (long 94.986115/lat 28.209435)">Pangin town</a> (28°12'33.96"N, 94°59'10.02"E, 450 m asl.), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=94.986115&amp;materialsCitation.latitude=28.209435" title="Search Plazi for locations around (long 94.986115/lat 28.209435)">East Siang district</a>, Arunachal Pradesh state, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=94.986115&amp;materialsCitation.latitude=28.209435" title="Search Plazi for locations around (long 94.986115/lat 28.209435)">Northeast</a> India, collected by members of the Systematics Lab, University of Delhi on 26 July 2009.</p><p>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 <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Sessa River</a>, nearby <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Sessa village</a> (27°6'4.02"N, 92°31'38.52"E, 1110 m asl.), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">West Kameng district</a>, Arunachal Pradesh state, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Northeast</a> India, collected by members of the Systematics Lab, University of Delhi on 07–09 August 2009 .</p><p>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 <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Sessa River</a>, nearby <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Sessa village</a> (27°6'4.02"N, 92°31'38.52"E, 1110 m asl.), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">West Kameng district</a>, Arunachal Pradesh state, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=92.52737&amp;materialsCitation.latitude=27.101116" title="Search Plazi for locations around (long 92.52737/lat 27.101116)">Northeast</a> India, collected by members of the Systematics Lab, University of Delhi on 07–09 August 2009 .</p><p>Provisionally referred specimens (see Remarks). One juvenile male (SDBDU 2009.132), and two juvenile females (SDBDU 2009.133 &amp; 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), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=96.40367&amp;materialsCitation.latitude=25.090334" title="Search Plazi for locations around (long 96.40367/lat 25.090334)">Hepu village</a>, 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 .</p><p>Holotype description (measurements in mm). Mature male (SVL 93.8) (Figures 22, &amp; 23A &amp; 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.</p><p>Forelimbs long, thin (Figures 22A &amp; B, &amp; 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&lt;II&lt;I&lt;III (FIL 13.7, FIIL 10.9, FIIIL 15.5, FIVL 10.3); interdigital webbing, subarticular and supernumerary tubercles absent; thenar and outer metacarpal tubercles weakly developed; finger tips flattened, not expanded relative to digit width, and with subcircular pads, terminal grooves on pads absent. Hindlimbs long, thin (Figures 22A &amp; B, &amp; 23A); thighs equal to shanks, and longer than feet (TL 47.4, SHL 50.2, FOL 45.8); toes long, dorsoventrally flattened, without lateral fringes (Figure 22E), relative toe lengths I&lt;II&lt;V&lt;III&lt;IV; toe tips flattened, not dilated, but with distinct oval-shaped pads, terminal grooves on pads absent; webbing rudimentary; inner metatarsal tubercles weakly defined, longitudinally ovalshaped; subarticular, supernumerary and outer metatarsal tubercles absent; ridge of callous tissue absent on ventral surface of toes.</p><p>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, “&gt;–&lt;” configuration, composed primarily of single row 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, flat, level with axilla on chest; femoral glands medium sized, 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, sparse on tympanic regions between tympanum and eyes, on dorsal surfaces of body and upper posterior surface of flanks, increasing in density posteriorly, present along all dorsal ridges, absent from all remaining surfaces.</p><p>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.</p><p>Refer to Figures 22 and 23A for further details of colouration and markings for holotype (in life, and in preservation).</p><p>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 IV&lt;II&lt;I&lt;III for most specimens, but IV=II&lt;I&lt;III observed on three (SDBDU 2009.1189, BNHS 6061, BNHS 6064); posterior edge of tongue varies from weakly, to moderately bifurcated, likely due to varying fixation conditions; dorsolateral ridges vary from weak to moderately well developed, always extending&gt;75% 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., “&gt;- (”, “&gt;––&lt;”, “&gt; &lt;”, “&gt; |”, “&gt;- &lt;”, or only V-shaped parietoscapular ridge present; coverage of dermal asperities varies mostly in density between individual males in comparison with holotype, however, on some specimens both black (or brown) and white asperities were observed, may sparsely cover upper lips, loreal region, front of snout, posterior dorsal surface of upper eyelids, head, and dorsal surface of tibia-tarsus joints; asperities white on female, restricted to dorsal surface of body, sparse on middorsum, increasing in density posteriorly to above cloaca; outer metacarpal tubercles not visible on most specimens; tubercle cover on flanks varies considerably, some with only sparse scattering of small tubercles, others with moderately dense cover of heterogeneous (large to small) sized tubercles (Figure 23 A–F). Dorsal and ventral markings vary considerably between individuals (see Figure 23 which represent extremes in variation).</p><p>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.</p><p>Morphological comparison. Characters used for comparing Megophrys periosa sp. nov. with its congeners do not include those from the provisionally assigned referred specimens.</p><p>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 II&lt;I in length, N =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%).</p><p>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).</p><p>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 &amp; 4; Appendix I, Table 3; Appendix II, Figures 1, 2 &amp; 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.</p><p>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.</p><p>Suggested common name: Giant Himalayan Horned Frog.</p><p>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).</p><p>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 &amp; 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 &amp; 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.</p><p>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 &amp; 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.</p></div>	https://treatment.plazi.org/id/03D6878AFFE4020DFF73FE9EFD51FF19	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
03D6878AFFE10270FF73FEF5FD74F92C.text	03D6878AFFE10270FF73FEF5FD74F92C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megophrys (Xenophrys) glandulosa Fei, Ye and Huang 1990	<div><p>Megophrys (Xenophrys) glandulosa Fei, Ye and Huang, 1990</p><p>(Figure 24; Table 1)</p><p>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].</p><p>Holotype. <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=100.44195&amp;materialsCitation.latitude=24.824167" title="Search Plazi for locations around (long 100.44195/lat 24.824167)">Adult</a> male (CIB 873112: Figure 24), from “Wuliang shan, Jingdong, Yunnan Province, altitude 1900m ” (= <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=100.44195&amp;materialsCitation.latitude=24.824167" title="Search Plazi for locations around (long 100.44195/lat 24.824167)">Wuliang Mountain</a> [~ 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).</p><p>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).</p><p>Examined specimens. Adult male (CIB 873112––photos only), holotype.</p><p>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), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=97.863884&amp;materialsCitation.latitude=27.293444" title="Search Plazi for locations around (long 97.863884/lat 27.293444)">Naung Mon township</a>, 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 .</p><p>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.</p><p>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.</p><p>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 IV&lt;I=II&lt;III (FIL 10.6, FIIL 10.6, FIIIL 13.1, FIVL 8.4); interdigital webbing, lateral fringes, subarticular and supernumerary tubercles absent; thenar tubercle distinct; metacarpal tubercles absent; digit tips slightly dilated, flattened, with subcircular pads, terminal grooves on pads absent. Hindlimbs relatively long, thin; thigh length slightly shorter than shanks, and longer than feet (TL 39.3, SHL 41.4, FOL 37.9); toes long, thin, relative toe lengths I&lt;II&lt;V&lt;III&lt;IV; digit tips not dilated but flattened, with subcircular pads, terminal grooves on pads absent; toes with thick basal webbing, rudimentary between digits I and II, remaining webbing formula as follows: II1.9– 3III 2.3–3.9IV3.8– 2V; moderately wide lateral fringes present on all toes; outer metatarsal, subarticular and supernumerary tubercles all absent; inner metatarsal tubercle present but barely distinguishable; ridge of thickened skin absent on ventral surface of digits.</p><p>Skin on dorsal surfaces of snout, head and back primarily smooth; throat, chest and limbs smooth; flanks with moderately large tubercles approximately forming two longitudinal rows, interspersed with densely scattered small granular tubercles; posterior thigh and cloacal region primarily smooth with small scattered weakly raised tubercles; tympanum mostly smooth, slightly concave; sides of head finely granular; palpebral horn absent, replaced by short fleshy bump; supratympanic ridge narrow anteriorly, extends from posterior orbital border, curves down along upper and posterior border of tympanum, considerably broadening posteriorly, terminating above shoulder; narrow, raised dorsolateral ridge extending posteriorly from behind supratympanic ridge to ~75% trunk length on both sides; weakly raised V-shaped parietoscapular ridge present, extending posteriorly from temporal region on each side, meeting medially at level of forelimb insertion; pectoral glands flat, level with axilla on chest; femoral glands slightly raised, on posterior surface of thighs, closer to knees than to cloaca; small black and white dermal asperities form a broad dense band circummarginally on gular region, moderately dense on tympanic regions between tympanum and eyes, and along upper lips, sparse on anterior dorsum of body restricted to dorsal ridges, increasing in density posteriorly across dorsum, absent from all remaining surfaces.</p><p>Colouration: 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.</p><p>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=I&lt;IV &lt;III, CAS 221395 has IV &lt;II&lt;I&lt;III; finger tips not dilated relative to finger width on CAS 221484 and CAS 221395; vomerine teeth absent on CAS 221484; dorsolateral ridges extend almost entire trunk length on CAS 221484; 5–10% of upper tympanum surfaces are concealed by supratympanic ridges; on CAS 221395, dermal asperities moderately dense on lateral surfaces of snout, dorsal surface of eyelids and dorsally on head adjacent to supratympanic ridges, sparse on dorsal tibia, outer and ventral tarsal surfaces; on CAS 221484, dermal asperities formed dense wide circummarginal band along lower jaw, sparse on tympanic regions (absent on tympanum) and on posterior lateral dorsum, absent on all remaining surfaces; V-shaped dorsal marking absent on CAS 221484 .</p><p>Secondary 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.</p><p>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).</p><p>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).</p><p>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).</p><p>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.</p><p>Suggested common name: Glandular White-lipped Horned Frog.</p><p>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).</p><p>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).</p><p>It is highly unlikely that the Bhutan (Wangyal 2013) and Indian (Meghalaya, Nagaland and West Bengal states: Ao et al. 2003; Deuti &amp; Ayyaswamy 2008; Mathew &amp; Sen 2010; Sangma &amp; 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.</p><p>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 .</p><p>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.</p></div>	https://treatment.plazi.org/id/03D6878AFFE10270FF73FEF5FD74F92C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mahony, Stephen;Kamei, Rachunliu G.;Teeling, Emma C.	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. Zootaxa 4523 (1): 1-96, DOI: 10.11646/zootaxa.4523.1.1
