taxonID	type	description	language	source
C83EE36BC447FD72FF1929C761FBFA65.taxon	description	(Figs. 2 A−F, 3 A−E)	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC447FD72FF1929C761FBFA65.taxon	materials_examined	Holotype MZB. Amph 22411 (field number ENS 14208; GenBank accession no. MT 710704; Figs. 2 A – B, 3 A – E; Megophrys sp. south in Fig. 1 and Table 1), an adult female collected from the eastern side of Bukit Barisan Mountain range in southern Sumatra, Lampung Province, Tanggamus Regency, Ulubelu District, Ngarip (5.280170 ° S, 104.557240 ° E, 1439 m a. s. l., Fig. 6), by Elijah Wostl, Kyle A. O’Connell and Ahmad Muammar Kadafi at 2040 h on 10 June 2013. Paratype UTA A- 66176 (formerly MZB. Amph 25988, field number ENS 17384; GenBank accession no. MT 710705; Fig. 2 C−D), subadult male collected at 1959 h on 8 July 2015 from the eastern side of Bukit Barisan Mountain range in southern Sumatra, Sumatra Selatan Province, Muara Enim Regency, Semendo Darat Ulu District, Segamit, near Gunung Patah (4.219890 ° S, 103.471250 ° E, 1624 m a. s. l., Fig. 6) by Elijah Wostl, Eric N. Smith, and Farid Akhsani. Referred specimens MZB. Amph 32593 (field number ENS 14202; GenBank accession no. MT 710706; Fig. 2 F) a juvenile male (SVLh 16.08 mm), same locality and collector as holotype; UTA A- 66177 (field number ENS 17403; GenBank accession no. MT 710707; Fig. 2 E) a juvenile male (SVLh 23.5 mm), same locality and collectors as paratype.	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC447FD72FF1929C761FBFA65.taxon	etymology	Etymology. The specific name selatanensis is derived from the Indonesian word Selatan (= south), as the new species exhibits a southern distribution within Sumatra and the Latin suffix – ensis meaning from that place. Suggested English common name. South-Sumatran Horned-Frog Suggested Indonesian name. Katak-tanduk Sumatra-selatan.	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC447FD72FF1929C761FBFA65.taxon	diagnosis	Diagnosis. The new species was assigned to the genus Megophrys based on the combination of the following morphological characters, as defined by Kuhl and van Hasselt (1822) and Delorme et al. (2006): (1) pointed snout profile bearing a pointed projection, protruding laterally beyond the lower jaw; (2) broad and flattened eyelid with a palpebral projection; (3) possession of a broad and depressed head; (4) conical spine at the corner of the mouth; (5) vertical pupil; (6) and presence of maxillary and vomerine teeth. Megophrys selatanensis sp. nov. can be diagnosed from geographically proximate congeners from the Sunda Shelf and the Philippines by the following combination of morphological characters: large body, stocky (SVLh 79.3 mm in adult female); snout acute with short rostral appendage (RSAL 0.3 % in adult female); relatively short triangular palpebral projection with acute tip (EHL 21.3 % UEWh in adult female); head wider than long (HW 126.4 % HLh in adult female); tympanum distinct, vertically elongated (TDH 52.4 % TDV in adult female); vomerine teeth present; a pair of dorsolateral folds extending from shoulder above axilla to groin; dorsal skin smooth with low and dense tuberculation; ventral skin smooth from throat to belly; short thigh (RTL 40.3 % in adult female), foot nearly as long as the thigh (FL 99.4 % TL in adult female); tibiotarsal articulation reaching posterior corner of eye; toes webbed at base.	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC447FD72FF1929C761FBFA65.taxon	description	Description of holotype (measurements in mm). Adult female, large body size (SVLh 79.3; SVL 79.1) and habitus stocky; head depressed and broad, wider (HW 38.1, 48.0 % SVLh) than long (HL 30.6, 38.6 % SVLh); snout short (SL 9.4, 11.9 % SVLh), pointed at tip, acute, with very short rostral appendage (SAL 0.3 % SVLh), laterally protruding and projecting beyond lower jaw; nostril positions laterally, closer to tip than to snout; eye positioned laterally, large, over three and a half times horizontal diameter of tympanum (ED 9.9, 365.1 % TDH), eye diameter (ED 9.9, 12.5 % SVLh) as long as snout-horn length (SLh 9.8, 12.3 % SVLh), about twice nostril-eye length (ED 210.6 % NEL), pupil vertical elliptical; canthus rostralis with sharp and angular ridge, lore sloping and concave; internarial distance (IND 7.6, 9.6 % SVLh) more than half of interorbital distance (IND 66.7 % IOD); palpebral projection length about 0.2 times of total upper eyelid width (EHL 1.8, 21.3 % UEWh), tip acute, surface smooth; upper eyelid edges with scattered small and low conical tubercles; tympanum distinct, smooth, oval, elongated vertically (TDH 2.7, 3.4 % SVLh; TDV 5.2, 6.5 % SVLh; TDH 52.4 % TDV); angular supratympanic fold, distinct, extending from behind eye, curving down around upper border of tympanum and ending above axilla; white conical tubercles present behind supratympanic fold and front of axilla; spinous gland on the corner of mouth on jaw angle; single row of maxillary teeth present; vomerine teeth present in two widely separated groups, at the level of posterior borders of choana; tongue lanceolate, notched posteriorly, without papillae. Forelimb slender, short, hand length about half of arm length (HAL 21.3, 55.9 % LAL), lower and upper arms slender; fingers moderately slender with rounded and swollen tips, unwebbed, lacking lateral fringes, finger length formula II = IV <I <III (fin 2 L 7.4, 9.3 % SVLh; fin 4 L 7.4, 9.3 % SVLh; fin 1 L 8.1, 10.2 % SVLh; and fin 3 L 10.0, 12.6 % SVLh); subarticular tubercles absent; outer palmar tubercle smaller than inner palmar tubercle (OPTL 3.9, 4.9 % SVLh; IPTL 7.2, 9.0 % SVLh). Hindlimb slender, moderately long (HLL 106.8, 134.7 % SVLh), thigh (FML 36.7, 46.3 % SVLh) slightly longer than tibia (TL 32.0, 40.1 % SVLh), about twice of tarsus length (TSL 17.6, 22.2 % SVLh), and nearly as long as foot (FL 31.8, 40.1 % SVLh); toe length formula I <II <V <III <IV (Toe 1 L 4.6, 5.8 % SVLh; Toe 2 L 5.6, 7.0 % SVLh; Toe 5 L 7.9, 9.9 % SVLh; Toe 3 L 8.9, 11.3 % SVLh; and Toe 4 L 17.5, 22.1 % SVLh); tibiotarsal articulation reaching to posterior corner of eye; inner metatarsal tubercle moderately large (IMTL 5.8, 7.4 % SVLh), outer absent; subarticular tubercles absent, toes with rounded and swollen tips, webbed only at base. Dorsal skin smooth, with low and dense tubercles, mostly on forelimbs, hindlimbs, and flanks; a pair of distinct dorsolateral folds, one on each side, extending from shoulder, above axilla to groin; vent without dorsal dermal accessory extension; three transverse folds on lower arm; five transverse folds on hindlimb, three on thigh and two on tibia; limbs skin laterally smooth, with small tubercles; ventral skin smooth, wrinkled in preservative; pair of white conical pectoral glands at base of axilla; a white conical femoral gland at mid flanks of posterior thigh. Coloration. In life (Fig. 2 A – B): pupil dark, iris golden-brown; base colour of dorsum uniform light brown; dorsal surface of forearm, hand, and hindlimbs slightly lighter than dorsum; upper limbs with dark brown transverse folds, three on forearm, three on thigh, and two on tarsus, fingers with dark brown crossbars, one on first and second fingers, two on third and fourth fingers; knee with irregular dark spots; dorsal surface and lateral sides of head light brown, with dark spots on upper and lower lips, from jaw angle to tip of snout; skin around eyes dark brown; flanks light brown, brighter than dorsum, unmarked; ventral surface light brown, with darker throat and chest; longitudinal markings over throat to pectoral region and dark blotches on belly; light brown blotches on ventral of limbs, with dark brown pattern near the joint between hand and lower arm. In preservative, dorsal surface darker, lateral and ventral surfaces fading to whitish, but pattern remains (Fig. 3 A – E). Variation. Type and referred specimens are morphometrically similar, despite large ontogenetic discrepancy between the adult female holotype (MZB. Amph 22411), subadult male paratype (UTA A- 66176), largest juvenile referred specimen (UTA A- 66177), and smaller juvenile referred specimen (MZB. Amph 32593). The adult specimen has slightly shorter rostral and palpebral projections, a shorter head, and shorter limbs, as compared to the subadult and juvenile specimens. The tibiotarsal joint, when the hindlimbs are adpressed forward, reaches the posterior corner of the eye in the adult and subadult, but only reaches the mid-eye in the juvenile. The morphometric variation is given in Table 2. In coloration, the dorsum of the subadult paratype (UTA A- 66176, Fig. 2 B) and the referred specimen (UTA A- 66177, Fig. 2 E) is brighter than that of the holotype, this is darker in the smaller referred specimen (MZB Amph 32593, Fig. 2 F). An irregular brighter brown pattern below the canthus rostralis is present in the paratype and the referred specimens but, it is absent in the holotype (Fig. 2 A, C, E – F). The paratype and referred specimens have a dark inverted triangle pattern on the parietal-orbital-scapular region. The paratype has orange-reddish coloration on the throat, chest, and ventral surface of the fingers and toes (Fig 2 C, D). The juvenile specimens (MZB Amph 32593 and UTA A- 66177) have darker brown throats and chests. Comparisons. Megophrys selatanensis sp. nov. is morphologically most similar to M. montana than to other geographically related congeners. Megophrys selatanensis sp. nov. has elongated dorsolateral folds that extend from the parietoscapular region to the groin, type I, as M. montana, M. parallela, M. lancip, M. nasuta, M. kalimantanensis Munir, Hamidy, Matsui, Iskandar, Sidik, and Shimada, and M. stejnegeri Taylor. Megophrys selatanensis sp. nov. differs from M. montana by having a shorter rostral appendage — SAL 0.3 − 0.6 % SVLh (vs. SAL reaching 3.5 % SVLh: Munir et al., 2018), shorter palpebral projection — EHL 21.3 – 24.3 % UEWh (vs. reaching 48.4 % UEWh: Munir et al., 2018), shorter ratio of foot to thigh — FL 96.6 − 99.4 % TL (vs. FL 106.2 − 115.6 % TL: Munir et al., 2018), and toes webbed only at base (vs. more developed web on second to fifth toes: Munir et al., 2018). Megophrys selatanensis sp. nov. differs from M. parallela by having elongated dorsolateral folds reaching the groin (vs. only reaching two-thirds of trunk), females being large — known adult female 79.1 mm SVL (vs. SVL ≤ 58.3 mm: Munir et al., 2018), a rostral appendage present (vs. absent: Inger and Iskandar, 2005; Munir et al., 2018), shorter ratio of foot to thigh — FL 96.6 – 99.4 % TL (vs. FL 105.4 – 112.5 % TL: Munir et al., 2018). The new species differs from M. lancip by having a less developed rostral appendage — SAL 0.3 − 0.6 % SVLh (vs. SAL 1.4 – 4.1 % SVLh: Munir et al., 2018), a shorter head length — HLh 38.0 – 38.5 % SVLh (vs. HLh 42.1 – 44.6 % SVLh: Munir et al., 2018), slightly wider head ratio to its length — HW 119.3 – 126.4 % HLh (vs. HW 103.2 – 116.0 % HLh: Munir et al., 2018), relatively shorter ratio of feet to thigh — FL 96.6 – 99.4 % TL (vs. FL 102.5 – 109.8 % TL: Munir et al., 2018), toes webbed only at base (vs. rather developed on third, fourth and fifth toes: Munir et al., 2018). Megophrys selatanensis sp. nov. differs from M. nasuta by the absence of additional lateral folds on flanks (vs. presence: Munir et al., 2018, 2019), shorter acute rostral appendage — SAL 0.3 − 0.6 % SVLh (vs. acuminate rostral appendage, SAL 1.8 − 9.2 % SVLh: Munir et al., 2019), shorter acute palpebral projection — EHL 2.2 − 3.4 % SVLh (vs. acuminate palpebral projection, EHL 4.4 − 13.0 % SVLh: Munir et al., 2019), shorter snout to nostril length — SNLh 3.7 − 5.1 % SVLh (vs. SNLh 5.1 − 13.8 % SVLh: Munir et al., 2019), shorter head length — HLh 38.0 − 40.5 % SVLh (vs. HLh 39.5 − 51.7. % SVLh: Munir et al., 2019), smaller ratio of palpebral projection to upper eyelid width — EHL 21.3 − 25.7 % UEWh (vs. EHL 32.7 − 61.4 % UEWh: Munir et al., 2019), and relatively wider head — HW 119.3 − 126.4 % HLh (vs. HW 92.9 − 119.5 % HLh: Munir et al., 2019). From M. kalimantanensis, the new species differs by the absence of additional lateral folds on flanks (vs. presence: Munir et al., 2019), female being smaller — known adult female 79.1 mm SVL (vs. 109.1 − 116.4 mm SVL: Munir et al., 2019), adpressed tibiotarsal articulation reaching posterior corner of eye in female (vs. posterior of tympanum, in female: Munir et al., 2019). From M. stejnegeri Taylor, Megophrys selatanensis sp. nov. differs by having the dorsolateral folds extending to the groin (vs. maximum of two-thirds length of trunk: Inger, 1954), female being slightly larger — known adult female 79.1 mm SVL (vs. SVL 57.0 – 77.8 mm: Taylor, 1920; Inger 1954), a rostral appendage present (vs. absent: Taylor, 1920; Inger, 1954), vomerine teeth present (vs. absent: Taylor, 1920; Inger, 1954), tympanum vertically elongated (vs. slightly rounded, see figure 37 by Diesmos et al., 2015 and figure 18 by Sanguila et al., 2016), and smooth skin on the temporal region (vs. tuberculate, see figure 37 by Diesmos et al., 2015 and figure 18 by Sanguila et al., 2016). Megophrys selatanensis sp. nov. is distinguished from M. kobayashii Malkmus and Matsui by having type I dorsolateral folds (vs. dorsolateral folds extended from parietoscapular region to mid-body, type II, see Table 3), additional lateral folds on flanks absent (vs. present: Malkmus and Matsui, 1997; Munir et al., 2018), female being smaller — known adult female 79.1 mm SVL (vs. SVL 99.0 – 109.0 mm: Malkmus and Matsui, 1997; Inger and Stuebing, 2005), a rostral appendage present (vs. absent: Malkmus and Matsui, 1997), and smaller and fewer tubercles on the body flanks (vs. larger and numerous: Malkmus and Matsui, 1997). From M. ligayae Taylor, the new species differs in having type I dorsolateral folds (vs. type II: Taylor, 1920; Inger, 1954; Munir et al., 2018), no additional lateral folds on flanks (vs. present: Taylor, 1920; Inger, 1954; Munir et al., 2018), female being smaller — known adult female 79.1 mm SVL (vs. SVL 90.0 mm: Inger, 1954), tibiotarsal articulation reaching posterior corner of eye (vs. tympanum: Taylor, 1920), and toes webbed only at base (vs. rather developed web on third, fourth, and fifth toes, see figure 37 C Diesmos et al., 2015). From M. edwardinae Inger, the new species differs by having type I dorsolateral folds (vs. absent: Inger, 1989), a rostral appendage present (vs. absent: Inger, 1989), vomerine teeth present (vs. absent: Inger, 1989), and smooth dorsal skin surface with low and dense tubercles (vs. dorsal surface with scattered round and elongated tubercles: Inger, 1989). Megophrys selatanensis sp. nov. can be distinguished from M. baluensis Boulenger by having type I dorsolateral folds (vs. dorsolateral folds formed by a series of elongated tubercle, type III: Inger and Stuebing, 2005: Munir et al., 2018), female being slightly larger — known adult female 79.1 mm SVL (vs. SVL ≤ 70 mm: Inger, 1966; Inger et al., 1995), a rostral appendage present (vs. absent: Boulenger, 1899; Inger, 1966; Inger et al., 1995), having a triangular palpebral projection (vs. small palpebral projection like tubercle: Inger, 1989; Inger et al., 1995), having smaller and fewer tubercles on body flanks (vs. larger and numerous: Inger et al., 2017), and adpressed tibiotarsal articulation reaching posterior of eye (vs. shoulder: Boulenger, 1899). Megophrys selatanensis sp. nov. differs from M. dringi Inger, Stuebing, and Tan, M. aceras Boulenger, and M. longipes Boulenger by the absence of Y, X,> – <or H shaped folds on dorsal (vs. presence: Boulenger, 1885, 1903; Inger et al., 1995, see Table 3). Furthermore, from M. dringi the new species being larger in female — known adult female 79.1 mm SVL (vs. SVL 55 mm: Inger et al., 1995; Inger and Stuebing, 2005), having a stocky body (vs. slender: Inger et al., 1995), a rostral appendage present (vs. absent: Inger et al., 1995), having a triangular palpebral projection (vs. small like tubercle: Inger et al., 1995), vomerine teeth present (vs. absent: Inger et al., 1995), distinct tympanum (vs. partially obscured: Inger et al., 1995). The new species differs from M. aceras by having a stocky body (vs. slender: Boulenger, 1903), a rostral appendage present (vs. absent: Boulenger, 1903), tibiotarsal articulation reaching to posterior corner of eye (vs. shoulder, angle of jaws or temporal area: Taylor, 1962), and toes webbed only at base (vs. rather possessing developed web between third, fourth, and fifth toes: Taylor, 1962; see figure 5 in Munir et al., 2018). From M. longipes, the new species differs being larger in female — known adult female 79.1 mm SVL (vs. SVL 49.0 – 65.0 mm: Inger et al., 1995), having a stocky body (vs. slender: Boulenger, 1885), a rostral appendage present (vs. absent: Boulenger, 1885), having a triangular palpebral projection (vs. small like tubercle: Boulenger, 1885; Taylor, 1962), shorter thigh — TL 0.41 − 0.44 SVL (vs. TL 0.54 – 0.65 SVL: Inger et al., 1995; Inger and Iskandar, 2005) and tibiotarsal articulation reaching posterior corner of eye (vs. far beyond tip of snout: Boulenger, 1885; Taylor, 1962).	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC447FD72FF1929C761FBFA65.taxon	distribution	Distribution and natural history. The holotype and one referred specimen (MZB Amph 32593) were collected in leaf litter on the edge of an old secondary forest in proximity to a coffee plantation and above a geothermal station in the province of Lampung, while the paratype (UTA A- 66176) and the other referred specimen (UTA A- 66177) were collected from an old secondary forest in the green corridor between the Gunung Patah and Gunung Agung. These locations are about 170 km apart along the eastern slope of the southern Barisan range of Sumatra. The tadpoles (ENS 14306 and ENS 14737) were collected from a small rocky stream in the old secondary forest near Lake Ranau, Lampung, 1108 – 1487 m a. s. l. (description of the tadpole of this new species is pending as we are preparing another manuscript to present detailed description of Sumatran Megophrys tadpoles). Most of the samples of M. selatanensis sp. nov. were collected from a small patched forest area in the eastern Bukit Barisan Selatan Mountain ranges, which is threatened by high deforestation. The exact distribution, population, breeding behavior, call, and other ecological information are unknown. The following anuran species have been found sympatrically with the new species, at Ngarip, Lampung: Indosylvirana nicobariensis (Stoliczka); Chalcorana chalconota (Schlegel); Wijayarana sumatrana (Yang); Hylarana erythraea (Schlegel); Ingerophrynus biporcatus (Gravenhorst); Leptophryne borbonica (Tschudi); Limnonectes kuhlii (Tschudi); L. macrodon (Duméril and Bibron); Limnonectes sp.; Microhyla gadjahmadai Atmaja, Hamidy, Arisuryanti, Matsui, Smith; M. palmipes Boulenger; Pelophryne sp.; Philautus larutensis (Boulenger); P. polymorphus Wostl, Riyanto, Hamidy, Kurniawan, Smith, and Harvey; P. thamyridion Wostl, Riyanto, Hamidy, Kurniawan, Smith, and Harvey; Phrynoides asper (Gravenhorst); Polypedates leucomystax (Gravenhorst); Rhacophorus achantharrhena Harvey, Pemberton, and Smith; R. catamitus Harvey, Pemberton, and Smith; and R. poecilonotus Boulenger; at the foothills of Gunung Patah, Sumatra Selatan: Wijayarana sumatrana; Limnonectes sp.; Microhyla sp.; Nyctixalus pictus (Peters); Philautus polymorphus; P. thamyridion; P. ventrimaculatus Wostl, Riyanto, Hamidy, Kurniawan, Smith, and Harvey; Phrynoides asper; Rhacophorus catamitus; and R. poecilonotus.	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC448FD68FF192A53600AFD76.taxon	description	(Figs. 4 A−D, 5 A−E)	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC448FD68FF192A53600AFD76.taxon	materials_examined	Holotype MZB. Amph 26098 (field number ENS 18664; GenBank accession no. MT 710708; Figs. 4 A−B, 5 A−E; Megophrys sp. north in Fig. 1 and Table 1), adult male collected at 2330 h on 5 August 2015 from Aceh Province, Aceh Tengah Regency, Linge District, Kute Robel (4.506444 ° N, 96.860750 ° E, 1638 m a. s. l), by Elijah Wostl, Ilham Fonna, and Muhammad Iksan (Fig. 6). Paratype UTA A – 66178 (field number ENS 21030; GenBank accession no. MT 710709; Fig. 4 C – D), a sub adult male collected on at 1856 h on 1 June 2016 from Aceh Province, Pidie Regency, Geumpang District, UPT V Geumpang (4.854540 ° N, 96.216540 ° E, 1086 m a. s. l) by Michael B. Harvey and Eric N. Smith (Fig. 6).	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC448FD68FF192A53600AFD76.taxon	etymology	Etymology. The specific name acehensis is derived from the province of Aceh in northern Sumatra and the Latin suffix – ensis meaning from that place. Suggested English common name. Aceh Horned Frog Suggested Indonesian name. Katak – tanduk Aceh	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC448FD68FF192A53600AFD76.taxon	diagnosis	Diagnosis. The new species was assigned to the genus Megophrys based on the combination of the following morphological characters, as defined by Kuhl and van Hasselt (1822) and Delorme et al. (2006): (1) pointed snout profile, bearing a pointed projection, protruding laterally beyond the lower jaw; (2) broad and flattened eyelid with palpebral projection; (3) possession of a broad and depressed head; (4) conical spine at the corner of mouth; (5) vertical pupil; (6) presence of maxillary and vomerine teeth. Megophrys acehensis sp. nov. can be diagnosed from its geographically relevant congeners in the Sunda Shelf and the Philippines by the following combination of characters: medium body size, stocky (SVLh 38.1 mm in adult male); snout pointed with a short, acute rostral appendage (RSAL 1.3 % in adult male); relatively short triangular palpebral projection with acute tip (EHL 25.1 % UEWh in adult male); head length relatively short (RHLh 37.2 % in adult male); head wider than its long (HW 113.6 % HLh in adult male); tympanum distinct, about one-third of eye diameter (TDH 35.6 % ED in adult male), nearly rounded (TDH 85.9 % TDV in adult male); vomerine teeth present; a pair of dorsolateral folds, extending from shoulder, above axilla to groin; dorsal and lateral skin tuberculate, shagreened on the throat to belly; short lower arm (RLAL 45.8 % in adult male); foot nearly as long as thigh (FL 96.6 % TL in adult male); tibiotarsal articulation reaching posterior of eye; toe webbing absent.	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC448FD68FF192A53600AFD76.taxon	description	Description of holotype (measurements in mm). Adult male, medium body size (SVLh 38.1, SVL 37.4) and habitus stocky; head depressed and broad, wider (HW 16.4, 42.9 % SVLh) than long (HL 13.3, 34.9 % SVLh); snout short (SL 4.2, 11.1 % SVLh), pointed at tip with acute short rostral appendage (SAL 0.7, 1.8 % SVLh), laterally protruding and projecting beyond lower jaw; nostril positioned laterally, near to snout than to eye; eye positioned laterally, large, nearly three times of tympanum horizontal diameter (ED 5.5, 280.5 % TDH), eye diameter slightly wider (ED 5.5, 14.4 % SVLh) than snout – horn length (SLh 5.4, 14.2 % SVLh), about two and three-quarter times of nostril – eye length (ED 275 % NEL), pupil vertical elliptical; canthus rostralis with sharp, angular ridge, lore sloping and concave; internarial distance (IND 4.0, 10.4 % SVLh) about two-thirds of interorbital distance (IND 68.4 % IOD); palpebral projection length about one-quarter of total upper eyelid width (EHL 1.2, 25.1 % UEWh), tip acute, surface smooth and scattered with small and low tubercles; tympanum distinct, smooth, oval, slightly rounded (TDV 2.3, 6.0 % SVLh; TDH 2.0, 5.1 % SVLh; TDH 85.9 % TDV); angular supratympanic fold, distinct, widened anteriorly, narrowed posteriorly, extending from behind eye, curving down around upper border of tympanum and ending above axilla; white conical tubercles behind the supratympanic fold and anterior to axilla; spinous gland on corner of mouth on jaw angle; single row of maxillary teeth present; vomerine teeth in two widely separated groups, at level posterior borders of choana; tongue lanceolate, notched posteriorly, without papillae; median subgular vocal sac present, having slit – like opening on each side of jaw commissures. * Fold forming Y, X or H on the parietoscapular region to the level of axilla; ** Dorsolateral fold shape: dorsolateral folds are elongated and extend from the parietoscapular region to the groin (Type I); dorsolateral folds extend from the central of parietoscapular region to mid-body (Type II); multiple dorsolateral folds - at least three or four - and they are discontinuous, formed by a series of elongated tubercles (Type III); dorsolateral folds are elongated and curve from the axillary region towards (and reaching) the posterior dorsal margin of tympanum (Type IV). Forelimb slender and short, hand length about half of arm length (HAL 9.5, 54.3 % LAL), lower arm proximally enlarged, wider than upper arm; fingers moderately slender, with rounded and swollen tips, unwebbed and lacking of lateral fringes; finger length formula I <II <IV <III (fin 1 L 2.6, 6.9 % SVLh; fin 2 L 3.2, 8.4 % SVLh; fin 4 L 3.5, 9.2 % SVLh; and fin 3 L 4.9, 12.8 % SVLh); subarticular tubercles absent; outer palmar tubercle smaller than inner (OPTL 1.9, 5.0 % SVLh; IPTL 2.5, 6.6 % SVLh). Hindlimb slender, moderately long (HLL 52.0, 136.5 % SVLh), thigh (FML 17.1, 44.8 % SVLh) slightly longer than tibia (TL 15.4, 40.4 % SVLh), about twice of tarsus length (TSL 8.5, 22.3 % SVLh), and nearly as long as foot (FL 14.9, 39.1 % SVLh); toe length formula I <II <V <III <IV (Toe 1 L 2.1, 5.4 % SVLh; Toe 2 L 3.3, 8.6 % SVLh; Toe 5 L 4.5, 11.8 % SVLh; Toe 3 L 4.6, 12.0 % SVLh; and Toe 4 L 7.8, 20.6 % SVLh); tibiotarsal articulation reaching to posterior corner of eye; inner metatarsal tubercle moderately large (IMTL 2.0, 5.1 % SVLh), outer absent; subarticular tubercles absent, toes with rounded and swollen tips, toes web absent. Dorsal skin surface tuberculate, low dense tubercles in entire dorsal and larger-sized tubercles mostly on forelimb, hindlimb, flanks and groin, shagreened in preservative; a pair of dorsolateral folds present, one on each side, extending from shoulder above the axilla to groin; vent with dorsal dermal accessory extension; transverse folds on limbs present, three on lower arm, three on thigh and three on tibia; limbs skin laterally tuberculate, larger size of tubercles than on dorsal side; ventral skin smooth shagreened on throat to chest, but smooth on belly and limbs, wrinkled in preservative; a pair of white conical pectoral glands, at base of axilla; white conical femoral glands, at the mid flanks of the posterior thigh; dark brown microspinules nuptial pads covering dorsal and median surfaces of the distal half of metacarpal and proximal half of basal phalanx of first and second fingers. Coloration. In life (Fig. 4 A – B, 5 A – B), pupil dark, iris golden – brown, base of dorsum immaculate dark brown with indistinct dark brown inverted triangle pattern on parieto-orbital to scapular region and inverted “ V ” like pattern on back, positioned closer to vent than to parietal region; dorsal surface of head, forelimb and hindlimb darker than lateral body; dark brown transverse folds present on dorsal surface of forearm, thigh and tibia, two on the forearm, three on thigh and three on tibia, fingers with dark brown crossbars, one on first and second fingers, two on third and fourth fingers; dorsomedial surfaces of first and second fingers with dark brown microspinular nuptial pads; knee with irregular dark brown spots; dorsal surface and lateral sides of head dark brown, irregular lighter brown pattern below canthus rostralis, light brown on underside of eyelids; body flanks dark brown and unmarked, lighter than dorsum; forelimb flanks dark brown, as dorsum; hindlimb flanks cream with dark brown blotches; ventral surface from throat to chest dark brown with heavy dark longitudinal marking; belly and underside of fore- and hindlimbs light cream with dark brown blotches; in preservative, pattern remains, but dorsal, lateral, and ventral surfaces are darker (Fig. 5 C). Variation. The two type specimens are morphometrically similar. The holotype has slightly longer rostral appendage, head, and snout-nostril and snout lengths, but a slightly shorter palpebral projection, and a thicker and narrower head. Morphometric variation is shown in Table 2. The transverse fold in the occipital region varies in degree of development, being distinct on the holotype but indistinct on the paratype. The dorsal coloration in life is dark brown on the holotype and orange-brown on the paratype. The ventral color of the two specimens is marbled with brown in both specimens but, the background color of the holotype is cream anteriorly and light grey posteriorly while that of the paratype is dark orange anteriorly and whitish posteriorly (Fig. 4 A−D). Comparisons. Megophrys acehensis sp. nov. differs from its most similar congener, M. parallela by having dorsolateral folds extending to the groin (vs. maximum of two-thirds length of trunk: Inger and Iskandar, 2005; Munir et al., 2018), a rostral appendage present (vs. absent: Inger and Iskandar, 2005; Munir et al., 2018), shagreened ventral surface from throat to belly (vs. smooth: Inger and Iskandar, 2005; Munir et al., 2018), and a relatively shorter foot to thigh ratio — FL 92.1 − 96.6 % TL (vs. 105.4 − 112.5 % TL: Munir et al., 2018). From M. montana, the new species differs by having tuberculate dorsal skin surface (vs. smooth: Munir et al., 2018), shagreened skin on throat (vs. smooth: Munir et al., 2018), a relatively short foot to thigh ratio — FL 92.1 − 96.6 % TL (vs. FL 106.2 − 115.6 % TL: Munir et al., 2018), and absent toe webbing (vs. rather developed webbing from second to fifth toes: Munir et al., 2018). From Megophrys selatanensis sp. nov., Megophrys acehensis sp. nov. differs by having a longer rostral appendage — SAL 1.3 – 1.8 % SVLh (vs. SAL 0.3 − 0.6 % SVLh), a slightly longer palpebral projection — EHL 25.1 – 28.4 % UEWh (vs. EHL 21.3 – 24.3 % UEWh), slightly narrower ratio of head width to its length — HW 114.1 − 115.3 % HLh (vs. HW 119.3 − 126.4 % HLh), wider ratio of tympanum to eye diameter — TDH 35.6 − 37.5 % ED (vs. TDH 27.3 − 28.5 % ED), tympanum nearly rounded — TDH 85.9 – 93.8 TDV (vs. vertically elongated, TDH 46.5 − 61.9 % TDV), slightly shorter lower arms — LAL 44.8 – 45.8 % SVLh (vs. LAL 48.0 – 49.2 % SVLh), tuberculate dorsal skin surface (vs. smooth), shagreened skin on throat (vs. smooth), and absent toe webbing (vs. toes webbed at base). Megophrys acehensis sp. nov. differs from M. lancip by males having a shorter rostral appendage — SAL 1.3 − 1.8 % SVLh (vs. SAL 2.9 − 3.1 % SVLh in males, but overlapped with females SAL 1.4 − 4.1 % SVLh: Munir et al., 2018), shorter head length — HLh 37.2 – 37.8 % SVLh (vs. HLh 42.1 – 44.6 % SVLh; Munir et al., 2018), wider head in males — HW 114.1 – 115.3 % HLh (vs. HW 103.2 – 105.8 % HLh in males, but overlapped with females HW 110.6 – 116.0 % HLh: Munir et al., 2018), wider tympanum relative to eye diameter — TDH 35.6 − 37.5 % ED (vs. TDH 25.7 − 28.0 ED: Munir et al., 2018), tympanum nearly rounded — TDH 85.9 – 93.8 TDV (vs. vertically elongated, TDH 61.1 − 69.1 % TDV), relatively short foot to thigh ratio — FL 92.1 − 96.6 % TL (vs. FL 102.5 − 109.8 % TL; Munir et al., 2018), and toe webbing absent (vs. rather developed on first to fifth toes, see Figs. 4 D, I, 5 B in Munir et al., 2018). Megophrys acehensis sp. nov. differs from M. nasuta by the absence of additional lateral flank folds (vs. presence: Munir et al., 2018, 2019), males being smaller — known adult male 37.4 mm SVL (vs. SVL 66.0 – 93.4 mm: Munir et al., 2019), shorter acute rostral appendage — SAL 1.3 − 1.8 % SVLh (vs. acuminate rostral appendage, SAL 1.8 − 9.2 % SVLh: Munir et al., 2019), and shorter acute palpebral projection — EHL 25.1 − 28.4 % UEWh (vs. acuminate palpebral projection, EHL 32.7 − 61.4 % UEWh: Munir et al., 2019). From M. kalimantanensis, the new species differs by the absence of additional flank folds (vs. present: Munir et al., 2019), smaller body size in males — known adult male 37.4 mm SVL (vs. SVL 64.3 – 100.6 mm: Munir et al., 2019), slightly longer rostral appendage — SAL 1.3 − 1.8 % SVLh (vs. SAL 0.1 − 0.8 % SVLh: Munir et al., 2019), and shorter palpebral projection — EHL 25.1 − 28.4 % UEWh (vs. 30.9 − 53.2 % UEWh: Munir et al., 2019). From M. stejnegeri, the new species differs by having dorsolateral folds extending to the groin (vs. maximum of two-thirds the length of the trunk: Inger 1954), males being smaller — known adult male 37.4 mm SVL (vs. adult male 40.6 − 60.0 mm: Taylor, 1920; Inger, 1954), vomerine teeth present (vs. absent: Taylor, 1920; Inger et al., 1954), and smooth skin on the posterior of tympanum (vs. tuberculate, see figure 37 by Diesmos et al., 2015 and figure 18 by Sanguila et al., 2016). Megophrys acehensis sp. nov. can be distinguished from M. kobayashii by having type I dorsolateral folds (vs. type II, see Table 3), absent additional lateral flank folds (vs. present: Malkmus and Matsui, 1997; Munir et al., 2018), males being smaller — known adult male 37.4 mm SVL (vs. 93.0 − 101.0 mm: Inger and Stuebing, 2005), rostral appendage present (vs. absent: Malkmus and Matsui, 1997), and smaller and fewer tubercles on flanks (vs. larger and numerous: Malkmus and Matsui, 1997). From M. ligayae, the new species differs by having type I dorsolateral folds, extended to the groin (vs. type II, reaching a maximum of two-thirds of trunk: Taylor, 1920; Inger, 1954; Munir et al., 2018), absent additional lateral folds (vs. present: Taylor, 1920; Inger, 1954; Munir et al., 2018), males being smaller — known adult male 37.4 mm SVL (vs. SVL 60.4 − 69.0 mm: Taylor, 1920; Inger, 1954), and toe webbing absent (vs. rather developed web on third, fourth, and fifth toes: Diesmos et al., 2015). Megophrys acehensis sp. nov. differs from M. edwardinae by having type I dorsolateral folds (vs. absent: Inger 1989), a rostral appendage present (vs. absent: Inger 1989), vomerine teeth present (vs. absent: Inger 1989), and slightly shorter thighs — TL 40.4 − 41.8 % SVLh (vs. TL 45.0 − 50.0 % SVL: Inger 1989). Megophrys acehensis sp. nov. can be distinguished from M. baluensis by having type I dorsolateral folds (vs. type III: Munir et al., 2018), males being slightly smaller — known adult male 37.4 mm SVL (vs. SVL 41.0 − 45.0 mm: Inger et al., 1995), a rostral appendage present (vs. absent: Boulenger, 1899; Inger, 1966; Inger et al., 1995), having a triangular palpebral projection (vs. small like tubercle: Inger, 1989; Inger et al., 1995), having smaller and fewer tubercles on body flanks (vs. larger and numerous: Inger et al., 2017), and adpressed tibiotarsal articulation reaching posterior of eye (vs. shoulder: Boulenger, 1899). Megophrys acehensis sp. nov. can be distinguished from M. dringi, M. aceras and M. longipes by the absence of Y, X,> – <or H shaped folds on dorsal (vs. presence: Boulenger, 1885, 1903; Inger et al., 1995, see Table 3). Furthermore, from M. dringi, the new species being smaller in male — known adult male 37.4 mm SVL (vs. SVL 43.0 − 47.0 mm: Inger et al., 1995), having a stocky body (vs. slender: Inger et al., 1995), a rostral appendage present (vs. absent: Inger et al., 1995), having a triangular palpebral projection (vs. small like tubercle: Inger et al., 1995), vomerine teeth present (vs. absent: Inger et al., 1995), distinct tympanum (vs. partially obscured: Inger et al., 1995). From M. aceras, the new species differs by having smaller body size in male — known adult male 37.4 mm SVL (vs. SVL 48.0 − 62.4 mm: Inger and Iskandar, 2005), a stocky body (vs. slender: Boulenger, 1903), rostral appendage present (vs. absent: Boulenger, 1903), tibiotarsal articulation reaching to posterior corner of eye (vs. shoulder, angle of jaws or temporal area: Taylor, 1962), toes web absent (vs. rather developed web on third, fourth, and fifth toes: Taylor, 1962, see figure 5 in Munir et al., 2018). From M. longipes, the new species differs by having a smaller body size in male — known adult male 37.4 mm SVL (vs. SVL 38.9 − 45.2 mm: Inger and Iskandar, 2005), having a stocky body (vs. slender: Boulenger, 1885), a rostral appendage present (vs. absent: Boulenger, 1885), having a triangular palpebral projection (vs. small like tubercle: Boulenger, 1885; Taylor, 1962), shorter thigh — TL 0.41 − 0.42 SVL (vs. TL 0.55 − 0.60 SVL: Inger and Iskandar, 2005) and tibiotarsal articulation reaching posterior corner of eye (vs. far beyond tip of snout; Boulenger 1885).	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
C83EE36BC448FD68FF192A53600AFD76.taxon	distribution	Distribution and Natural History. The holotype of Megophrys acehensis sp. nov. was collected on leaf litter in a sloping area at the edge of a primary forest near a stream, while the paratype was collected from leaf litter in a palm oil plantation near the edge of an old secondary forest. Landslides, new road development, and monoculture forests have become major threats at the holotype locality, while the threats at the paratype locality were the land use changes and water pollution from palm oil fields. The precise distribution, population, habitat requirements, breeding behavior, call and tadpole information are unknown. The following anuran species have been found sympatrically with the new species, at holotype locality: Limnonectes sp; Philautus larutensis; Sumaterana dabulescens Arifin, Smart, Hertwig, Smith, Iskandar, and Hass; at the paratype locality: Chalcorana chalconota; Leptophryne borbonica; Limnonectes kuhlii; L. macrodon; Limnonectes sp.; Rhacophorus catamitus; Philautus sp. and Pulchrana fantastica Arifin, Cahyadi, Smart, Jankowski, and Haas.	en	Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir, Smith, Eric N. (2021): Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia. Zootaxa 5057 (4): 503-529, DOI: https://doi.org/10.11646/zootaxa.5057.4.3
