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Zhang, Yinpeng, Rowley, Jodi J. L., Liu, Xiaolong, Nguyen, Tao Thien, Nguyen, Huy Quoc, Tapley, Benjamin, Nguyen, Luan Thanh, Yang, Yun, Cutajar, Timothy, Zhang, Ying, Yuan, Zhiyong (2021): Amolops caelumnoctis Rao & Wilkinson, 2007, a Junior Synonym of A. splendissimus Orlov & Ho, 2007 (Amphibia: Anura: Ranidae). Zootaxa 5057 (2): 181-200, DOI: https://doi.org/10.11646/zootaxa.5057.2.2

Tapley, Benjamin, Cutajar, Timothy, Nguyen, Luan Thanh, Portway, Christopher, Mahony, Stephen, Nguyen, Chung Thanh, Harding, Luke, Luong, Hao Van, Rowley, Jodi J. L. (2021): A new potentially Endangered species of Megophrys (Amphibia: Megophryidae) from Mount Ky Quan San, north-west Vietnam. Journal of Natural History (J. Nat. Hist.) 54 (39-40): 2543-2575, DOI: 10.1080/00222933.2020.1856952, URL: http://dx.doi.org/10.1080/00222933.2020.1856952

Megophrys (Xenophrys) maosonensis Figs. 3E and 8 Molecular data: A 16S sequence was generated from a tissue sample obtained from the single tadpole (VNMN010904). Uncorrected p -distance between the tadpole in this study and a M. maosonensis specimen also collected in the rough proximity of one of the type specimens; Sa Pa District, Lao Cai Province, Vietnam (GenBank accession number KX 811786) was 0.0 %. Collection site: The following tadpole description is based on two specimens at Gosner stage 25 (VNMN010904 and HLNP 2018090900015). The specimens were found feeding in a 8 m wide stream in disturbed evergreen forest on Mount Fansipan, Hoang Lien National Park, Cat Cat river, Sa Pa District, Lao Cai Province, Vietnam (22.3214��N 103.8264��E, 1244 m asl; Figs. 1 and 2D), on 9 September 2018 by Luan Nguyen, Chung Nguyen and Luong Hoang. Morphology: The body is elongated and slender; the nares are oval and are approximately equal in distance from the eyes and the snout, the rims of the nares are serrated and raised from the body wall, the internarial distance is less than the interorbital distance; the eyes are positioned dorsolaterally, the pupils are round; the spiracle is sinistral and the spiracular tube protrudes from one third of the way along the body, anterior to maximal trunk width and opens laterally; the tail length is 76.5���77.9% of the total length; the dorsal tail fin inserts behind the body-tail junction, the dorsal fin is low, particularly towards the proximal half of the tail length; the basal tail width is 51.2���61.0%, of the maximal trunk width; the tail tip is narrowly rounded; the oral disc is subterminal and antero-dorsal; the lateral corners fold medially and turn upward when not fully extended (and in preservative), the mean width of the umbelliform oral disc makes up 63.4���68.3% of the maximal width of the trunk; in life, the maximal BW is 73.2% (n =1)the maximal width of the fully extended ODW and the width of the umbelliform oral disc exceeds the maximal width of the trunk; the lower lip is bi-triangular shaped; marginal papillae are absent; five rows of submarginal papillae are present on the upper lip when they are counted medially at the maximal width of the oral disc; these are longitudinally oblong shaped and become substantially smaller in the outermost row; four rows of longitudinally oblong submarginal papillae are present on the lower lip when they are counted medially at the maximal width of the oral disc; these are regularly positioned and become smaller on the outermost row the lower lip is deeper than the upper lip; labial teeth are absent; the upper jaw sheath is serrated and has a medial notch; the lower jaw sheath lacks a medial notch. See Table 1 for measurements. Colour in life: (Fig. 8) Dorsally, the head and body are brown with dark brown speckles which become increasingly dense towards the posterior half of the body; a dark brown bar runs from each nare to the anterior edge of the eye; there are obvious pale neuromasts; the oral disc is translucent orange brown with darker brown submarginal papillae; the dorsal apex of the tail muscle on the anterior half of the tail is orange brown and forms a saddle which is speckled with dark brown, the lower margins of the orange brown saddle fade to cream bordered by a dark brown band which runs half way along the tail length on the lateral surface; at the point where the tail meets the body there is an area of dense dark brown flecks which form a thin line which extends from the body-tail junction and runs along the midline of the lateral tail surface; the dorsal and ventral tail fins are opaque, beige brown with a few dark brown speckles; the ventral surface is a translucent cream with and speckled with metallic grey blue; the lower lip is bordered by a ���V��� shaped area of dark stippling when viewed from the ventral surface; the sclera of the eye is broad and black with green-gold flecks; the pupil is black and round; the iris orange and speckled with black dots. Colour in preservative: (Fig. 3E) The body is grey with darker speckles; the oral disc is a translucent grey with dark brown submarginal papillae; the dorsal and ventral fins are opaque, pale grey with few visible darker speckles; the ventral surface is dark grey. Variation: The tadpole of M. maosonensis at Stages 37���39 was described previously by Fei et al. (2009; see Table 1 for measurements) as M. major Boulenger; a species now only definitively known from northeast India (Mahony et al. 2018), although species identity was not apparently supported with molecular data in Fei et al. (2009). The line drawing depicts a tadpole with a very pointed tail tip, the specimens from the Hoang Lien Range have a narrowly rounded tail tip (Fig. 8). Robust comparison cannot be made with the description of Fei et al. (2009) as the tadpoles were of a much later stage (37���39) than those collected in the Hoang Lien Range (Stage 25)., Published as part of Tapley, Benjamin, Nguyen, Luan Thanh, Cutajar, Timothy, Nguyen, Chung Thanh, Portway, Christopher, Luong, Hao Van & Rowley, Jodi J. L., 2020, The tadpoles of five Megophrys Horned frogs (Amphibia: Megophryidae) from the Hoang Lien Range, Vietnam, pp. 35-52 in Zootaxa 4845 (1) on pages 48-49, DOI: 10.11646/zootaxa.4845.1.3, http://zenodo.org/record/4477449, {"references":["Fei, L., Hu, S. Q., Ye, C. Y. & Huang, Y. Z. (2009) Fauna Sinica. Amphibia. Volume 2. Anura. Chinese Academy of Science, Science, Beijing, XIII + 957 pp.","Mahony, S., Kamei, R. G., Teeling, E. C. & Biju, S. D. (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 - 96."]}

Megophrys (Panophrys) fansipanensis Figs. 3B and 5 Molecular data: 16S sequences were generated from tissue samples collected from two tadpoles (VNMN010901 & HLNP 20171230 00011) and uncorrected p -distance between these sequences and that of the holotype of Megophrys fansipanensis also collected on Mount Fansipan, Sa Pa District, Lao Cai Province, Vietnam (GenBank accession number MH 514886) was 0.0 %. Collection site: The following tadpole description is based on two specimens (VNMN010901 & HLNP 20171230 00011) at Stage 35. Specimens were collected from a small pool in a 1 m wide low-gradient stream in disturbed evergreen forest on Mount Fansipan, Hoang Lien National Park (HLNP), Sa Pa District, Lao Cai Province, Vietnam (22.3281��N 103.7817��E, 2242 m asl; Figs. 1 and 2C), collected on 30 December 2017 by Luan Nguyen and Chung Nguyen. Morphology: The body is elongated and slender; the nares are oval and their rims raised from the body wall, the nares are closer to the eyes than to the snout; the internarial distance is less than the interorbital distance; the eyes are positioned dorsolaterally, the pupils are round; the spiracle is sinistral and the spiracular tube protrudes from midway along the body wall just anterior to the maximal trunk width and opens laterally; the tail length ranges from 69.1���72.1% (n =2) of the total length; the dorsal tail fin inserts behind the body-tail junction, the dorsal fin is low, particularly towards the proximal half of the tail length; the basal tail width ranges from 53.1���60.3%, (n =2) of the maximal trunk width; the tail tip is pointed; the oral disc is subterminal and antero-dorsal; the lateral corners fold medially and turn upward when not fully extended (and in preservative), the width of the umbelliform oral disc makes up 74.1���82.9%, (n =2) of the maximal width of the trunk; in life the maximal BW is 64.8% (n =1) the maximal width of the fully extended ODW; the lower lip is bi-triangular shaped when fully expanded in live specimens; marginal papillae are absent; five rows of submarginal papillae are present on the upper lip when they are counted medially at the maximal width of the oral disc; these are regularly positioned and rounded, and become substantially smaller in the outermost row; five rows of rounded submarginal papillae are present on the lower lip when they are counted medially at the maximal width of the oral disc; these are regularly positioned, longitudinally oblong shaped at the edge of the disc and become increasingly rounded towards the point at which the maximal width of the disc is at its greatest, the submarginal papillae become smaller and more longitudinally elongated in shape on the outermost row; the lower lip is deeper than the upper lip; labial teeth are absent; the upper jaw sheath is serrated and has a medial notch; the lower jaw sheath lacks a medial notch. See Table 1 for measurements. Colour in life: (Fig. 5) Dorsally, the head and body are brown with dark brown speckles which become increasingly dense towards the posterior half of the body; there are obvious pale neuromasts; the oral disc is a translucent beige brown with dark brown submarginal papillae; the dorsal apex of the tail has a light brown saddle, the margins of which are bordered by a dark brown band on the lateral surfaces of the tail, this saddle extends from the body and terminates half way along the length of the tail; at the point where the tail meets the body there is a brown stripe that extends from the body-tail junction and runs along the midline of the lateral tail surface, this stripe becomes increasingly broken and fades into brown blotches half way along the tail; the dorsal and ventral tail fins are opaque, pale yellowish brown with a few dark brown speckles; the venter is a translucent grey brown and speckled with metallic blue and flecked with dark brown; the gills and coils of the gut are visible in the ventral view through the transparent ventral skin; the sclera of the eye is broad and black with green-gold flecks; the pupil is black and rounded; the iris is orange and speckled with black dots; the outer margin of the upper jaw sheath is dark brown. Colour in preservative: (Fig. 3B) The body is brown with darker speckles; the oral disc is a translucent grey brown with dark brown submarginal papillae; the dorsal and ventral fins are opaque, pale grey brown with darker speckles; the venter is speckled grey and brown. Variation: HLNP 20171230 00011 is larger than VNMN010901 (Fig. 5) The ventral surface of the body of VNMN010901 is more densely speckled with metallic blue. TMW/BW is greater in HLNP 20171230 00011., Published as part of Tapley, Benjamin, Nguyen, Luan Thanh, Cutajar, Timothy, Nguyen, Chung Thanh, Portway, Christopher, Luong, Hao Van & Rowley, Jodi J. L., 2020, The tadpoles of five Megophrys Horned frogs (Amphibia: Megophryidae) from the Hoang Lien Range, Vietnam, pp. 35-52 in Zootaxa 4845 (1) on page 44, DOI: 10.11646/zootaxa.4845.1.3, http://zenodo.org/record/4477449

Megophrys (Atympanophrys) gigantica Figs. 1, 2A, 3A, and 4 Molecular data: 16S sequences were generated from tissue samples of two tadpoles. Uncorrected p -distance between the tadpoles in this study and sequences from M. gigantica specimens collected approximately 390 km to the northwest in the proximity of the type locality; Mount Wuliang, Yunnan Province, China (GenBank accession numbers; MH 406775; MH 406776 & KX 811898) were 0.2 %. Collection site: The following tadpole description is based on 5 specimens VNMN010898, VNMN010899, VNMN010900, HLNP 20180320 00009 & HLNP 20180320 00010) at Stage 35. Specimens were found actively feeding near the bank in a large pool in a 2 m wide stream in heavily disturbed evergreen forest on Mount Pu Ta Leng, Bat Xat Nature Reserve, Bat Xat District, Lao Cai Province, Vietnam (22.4322��N 103.6297��E, 1912 m asl; Figs. 1 and 2A), collected at 15:00 h on 20 March 2018 by Luan Nguyen, Christopher Portway, Chung Nguyen and Benjamin Tapley. Morphology: The body is longitudinally oval shaped and dorsally compressed; the nares are oval and are closer to the eyes than to the snout, the rims of the nares are serrated, raised from the body wall and open anterolaterally; the internarial distance is less than the interorbital distance; the eyes are positioned dorsolaterally, the pupils are round; the spiracle is sinistral and the spiracular tube protrudes from nearly two thirds of the way along the body wall just anterior to the maximal trunk width, the spiracular tube opens laterally; the mean tail length is 66.6% (63.2���68.4%, n =5) of the total length; the dorsal tail fin inserts behind the body-tail junction, the dorsal fin is low, particularly at the proximal half of the tail length; the mean basal tail width is 42.1% (37.2���45.1%, n =5) of the maximal trunk width, the tail tip is broadly rounded; the oral disc is subterminal and antero-dorsal, the lateral corners fold medially and turn upward when not fully extended (and in preservative), the mean width of the umbelliform oral disc makes up 92.4% (82.4���96.7%, n =5) of the maximal width of the trunk; in life the maximal BW is 62.6% the maximal width of the fully extended ODW (n =1); the lower lip is triangular with basal lobes when the mouthparts are fully expanded in life, we define this as hastate shaped (Fig. 4B), marginal papillae are absent; six rows of submarginal papillae are present on the upper lip when they are counted medially at the maximal width of the oral disc; these are regularly positioned; those present medially at the maximal width of the oral disc are rounded, but those in the innermost two rows become increasingly oblong shaped towards the edges of the disc; five rows of rounded submarginal papillae are present on the lower lip when they are counted medially at the maximal width of the oral disc; these are regularly positioned, longitudinally oblong shaped on the three inner rows and they become smaller and circular in shape on the outermost row; the lower lip is deeper than the upper lip; labial teeth are absent; the upper jaw sheath is serrated and has a deep medial notch; the lower jaw sheath is serrated and lacks a medial notch. See Table 1 for measurements. Colour in life: (Fig. 4) Dorsally, the head and body are light brown with pale neuromasts; the area surrounding the nares is dark brown; the oral disc is a translucent tan brown with dark brown submarginal papillae; the dorsal and ventral fins are opaque, pale yellowish brown without speckles; the lower fin is more yellow in colour than the upper fin and also without speckles; the ventral and lateral sides of the body are a translucent dark grey and speckled with white; the underside of the oral disc is speckled with white; the gills and coils of the gut are visible in the ventral view through the transparent ventral skin; the sclera of the eye is black with green-gold flecks; the pupil is black and rounded; the iris is orange and speckled with black dots; the outer margin of the upper jaw sheath is dark brown. Colour in preservative: (Fig. 3A) Body brown; the gills and coils of the gut are visible in the ventral view through the transparent ventral skin; the oral disc is translucent grey brown with darker brown submarginal papillae; the dorsal and ventral fins are an opaque, pale grey brown; the venter is grey; the iris is dark grey. Variation: VNMN010899 has a shorter tail than the other M. gigantica specimens collected from the Hoang Lien Range, this is probably due to a historic tail injury and subsequent regeneration (Figs. 3A and 4A). The tadpole of M. gigantica at Stage 34 was described previously by Fei et al. (2009) although it is not clear if species identity was supported with molecular data. The description is broadly congruent with the Stage 35 specimens collected in this study but there are some notable differences.At stage 34 the TTL is reported as 39.0 mm (Fei et al. 2009); this is considerably smaller than the mean TTL of 50.7 mm in this study. Fei et al. (2009) describe the body and tail muscle as having grey patterns (also depicted in the associated line drawing). The specimens we collected in the Hoang Lien Range lacked grey patterns on the body and on the tail muscle., Published as part of Tapley, Benjamin, Nguyen, Luan Thanh, Cutajar, Timothy, Nguyen, Chung Thanh, Portway, Christopher, Luong, Hao Van & Rowley, Jodi J. L., 2020, The tadpoles of five Megophrys Horned frogs (Amphibia: Megophryidae) from the Hoang Lien Range, Vietnam, pp. 35-52 in Zootaxa 4845 (1) on pages 37-39, DOI: 10.11646/zootaxa.4845.1.3, http://zenodo.org/record/4477449, {"references":["Fei, L., Hu, S. Q., Ye, C. Y. & Huang, Y. Z. (2009) Fauna Sinica. Amphibia. Volume 2. Anura. Chinese Academy of Science, Science, Beijing, XIII + 957 pp."]}

Megophrys (Panophrys) hoanglienensis Figs. 3D and 7 Molecular data: A 16S sequence was generated from tissue samples collected from one tadpole (VNMN010903) and uncorrected p -distance between this sequence and that of the holotype of M. hoanglienensis also collected on Mount Fansipan, Sa Pa District, Lao Cai Province, Vietnam (Genbank accession number MH 514889) was 0.0 %. Collection site: The following tadpole description is based on a single specimen (VNMN010903) at Stage 26. The specimen was found feeding in a 5 m wide stream in disturbed evergreen forest on Mount Fansipan, Hoang Lien National Park, Tam Duong District, Lai Chau Province, Vietnam (22.3483��N 103.7700��E, 1901 m asl; Figs. 1 and 2B), on 10 September 2018 by Luan Nguyen, Chung Nguyen and Luong Hoang. Morphology: The body is longitudinally oval shaped and dorsally compressed; the nares are oval and are closer to the eyes than to the snout, the rims of the nares are serrated and raised from the body wall, the internarial distance is less than the interorbital distance; the eyes are positioned dorsolaterally, the pupils are round; the spiracle is sinistral and the spiracular tube protrudes from midway along the body wall and anterior to maximal trunk width and opens laterally; the tail length is 73.2% of the total length; the dorsal tail fin inserts behind the body-tail junction, the dorsal fin is low, particularly towards the anterior half of the tail length; the basal tail width is 43.5%, of the maximal trunk width, the tail tip is pointed; the oral disc is subterminal and antero-dorsal; the lateral corners fold medially and turn upward when not fully extended (and in preservative), the mean width of the umbelliform oral disc makes up 47.8% of the maximal width of the trunk, in life the width of the umbelliform oral disc exceeds the maximal width of the trunk; in life the maximal BW is 79.3% (n =1) the maximal width of the fully extended ODW; the lower lip is bi-triangular shaped; marginal papillae are absent, the edges of the lips are smooth; four rows of submarginal papillae are present on the upper lip when they are counted medially at the maximal width of the oral disc; these are longitudinally oblong shaped at the edge of the disc and become increasingly rounded towards the maximal width of the disc, and become substantially smaller in the outermost row; five rows of longitudinally oblong submarginal papillae are present on the lower lip when they are counted medially at the maximal width of the oral disc; these are regularly positioned and become smaller on the outermost row; the lower lip is deeper than the upper lip; labial teeth are absent; the upper jaw sheath is serrated and has a medial notch; the lower jaw sheath lacks a medial notch. See Table 1 for measurements. Colour in life: (Fig. 7) Dorsally, the head and body are dark brown, a darker brown irregular shaped bar runs dorsally from nare to nare, a large reddish brown blotch is present posterior to each of the eyes, the area between these reddish brown blotches and mid dorsum is reticulated with blackish brown; the upper lateral surface of the body is beige with darker brown reticulations and light grey flecks, the skin of the lower lateral surface of the body is also flecked with grey over slightly translucent skin, the colour of the organ structures is visible beneath; the neuromasts are distinct; the oral disc is translucent beige brown, the colouration becoming darker towards the outer margins of the disc, the submarginal papillae are dark brown; the dorsal apex of the tail has light brown saddle, the margins of which are bordered by a dark brown band on the lateral surfaces of the tail, this saddle extends from the body and terminates closer to body than to tail tip; at the point where the tail meets the body there is a brown stripe that extends from the body-tail junction and runs along the midline of the lateral tail surface, this stripe becomes increasingly broken and fades into brown blotches half way along the length of the tail; the dorsal and ventral tail fins are opaque, the upper tail fin is a pale yellowish brown with many dark brown speckles, the lower tail fin is largely translucent with many dark brown speckles; the venter is translucent, speckled with metallic grey blue flecks; the sclera of the eye is broad and black with green-gold flecks; the pupil is black and round; the iris is orange and speckled with black dots. Colour in preservative: (Fig. 3D) The body is brown with darker speckles; the oral disc is a translucent grey brown with dark brown submarginal papillae; the dorsal and ventral fins are opaque, upper fin pale grey brown with darker speckles; the venter is speckled grey and brown., Published as part of Tapley, Benjamin, Nguyen, Luan Thanh, Cutajar, Timothy, Nguyen, Chung Thanh, Portway, Christopher, Luong, Hao Van & Rowley, Jodi J. L., 2020, The tadpoles of five Megophrys Horned frogs (Amphibia: Megophryidae) from the Hoang Lien Range, Vietnam, pp. 35-52 in Zootaxa 4845 (1) on pages 46-47, DOI: 10.11646/zootaxa.4845.1.3, http://zenodo.org/record/4477449

Megophrys (Panophrys) jingdongensis Figs. 3C and 6 Molecular data: A 16S sequence was generated from a tissue sample obtained from the single tadpole (VNMN010902). Uncorrected p -distance between the tadpole in this study and a M. jingdongensis specimen collected approximately 390 km to the northwest in the proximity of the type locality; Wenlong, Jingdong County, Yunnan Province, China (GenBank accession number KX 811874) was 0.1 %. Collection site: The following tadpole description is based on a single specimen (VNMN010902) at Stage 25. The specimen was found near the bank in a large, 2 m wide pool in heavily disturbed evergreen forest on Mount Pu Ta Leng, Bat Xat Nature Reserve, Bat Xat District, Lao Cai Province, Vietnam (22.4322��N 103.6297��E, 1912 m asl; Figs. 1 and 2A), at 15:00 h on 20 March 2018 by Luan Nguyen, Christopher Portway, Chung Nguyen and Benjamin Tapley. Morphology: The body is elongated and slender; the nares are oval and are closer to the eyes than to the snout, the rims of the nares are serrated and raised from the body wall, the internarial distance is less than the interorbital distance; the eyes are positioned dorsolaterally, the pupils are round; the spiracle is sinistral and the spiracular tube protrudes from midway along the body wall and anterior to maximal trunk width and opens laterally; the tail length is 68.1% of the total length; the dorsal tail fin inserts behind the body-tail junction, the dorsal fin is low, particularly towards the proximal half of the tail length; the basal tail width is 48.8% of the maximal trunk width, the tail tip is rounded; the oral disc is subterminal and antero-dorsal; the lateral corners fold medially and turn upward when not fully extended (and in preservative), the width of the umbelliform oral disc makes up 63.4% of the maximal width of the trunk; in life the maximal BW is 80.4% (n =1) the maximal width of the fully extended ODW; the lower lip is bi-triangular shaped; marginal papillae are absent; regularly positioned, round and oblong shaped submarginal papillae are present on both the upper and lower lip; the lower lip is deeper than the upper lip; labial teeth are absent; the upper jaw sheath is serrated and has a medial notch; the lower jaw sheath lacks a medial notch. See Table 1 for measurements. Colour in life: (Fig. 6) Dorsally, the head and body are dark brown with a series of cream blotches bordered by orange flecks, diffuse beige flecks cover the dorsal and lateral surfaces of the body; neuromasts are indistinct; the oral disc is a translucent beige brown and the submarginal papillae are dark brown; the dorsal apex of the tail has four cream saddles, the margins of which are bordered with dark brown, the saddles extend from the body and terminate half way along the length of the tail; at the point where the tail meets the body, there is a brown stripe that extends from the body-tail junction and runs along the midline of the lateral tail surface, this stripe becomes increasingly broken and fades into brown blotches half way along the length of the tail, the dorsal and ventral tail fins are opaque, the dorsal fin has many dark brown speckles along its length; the ventral fin has few, very dispersed dark brown speckles; the vent is grey brown and speckled with metallic blue; the sclera of the eye is black; the pupil is black and round; the iris is gold, speckled with black dots; the gills and coils of the gut are visible in the ventral view through the transparent ventral skin. Colour in preservative: (Fig. 3C) The body is brown with lighter speckles and blotches; the oral disc is a translucent grey brown with dark brown submarginal papillae; the dorsal and ventral fins are opaque, pale grey brown with darker speckles; the venter is speckled grey and brown. Variation: The tadpole of M. jingdongensis at Stage 37 was described previously by Fei et al. (2009) although it is unclear if species identity was supported with molecular data. The description is broadly congruent with the specimen collected in this study but there are some notable differences. In the text of Fei et al. (2009) the tail tip is described as blunt, but the line drawing of the tadpole shows a specimen with a pointed tail tip. Fei et al. (2009) describe the body of M. jingdongensis as Stage 37 as unmarked, this is incongruent with our observation of a single specimen at Stage 25 where the body is heavily marked both in life and in preservative., Published as part of Tapley, Benjamin, Nguyen, Luan Thanh, Cutajar, Timothy, Nguyen, Chung Thanh, Portway, Christopher, Luong, Hao Van & Rowley, Jodi J. L., 2020, The tadpoles of five Megophrys Horned frogs (Amphibia: Megophryidae) from the Hoang Lien Range, Vietnam, pp. 35-52 in Zootaxa 4845 (1) on pages 44-46, DOI: 10.11646/zootaxa.4845.1.3, http://zenodo.org/record/4477449, {"references":["Fei, L., Hu, S. Q., Ye, C. Y. & Huang, Y. Z. (2009) Fauna Sinica. Amphibia. Volume 2. Anura. Chinese Academy of Science, Science, Beijing, XIII + 957 pp."]}

Megophrys (Panophrys) fansipanensis sp. nov. Figs. 1, 3���7 & Table 2. Holotype: Adult male (VNMN 2018.01), found perched on a stick 3 m above a 2 m wide low-gradient stream in disturbed evergreen forest on Mount Fansipan, Hoang Lien National Park (HLNP), Sa Pa District, Lao Cai Province, Vietnam (22.32827�� N, 103.78184�� E, 2242 m asl; Fig. 1), collected at 19:15 h on 12 June 2012 by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. Paratypes: Adult female (VNMN 07033), found resting on leaf litter associated with bamboo forest beside a 1 m wide cascading stream on Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam (22.31458�� N, 103.76574�� E, 2783 m asl; Fig 1), collected at 20:00 h on 0 9 June 2012; four adult males (AMS R186105���08), found in the vicinity of a 1 m wide cascading stream on Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam (22.31349�� N, 103.76521�� E, 2813 m asl; Fig. 1), on 10 June 2012; four adult males (AMS R186109���12), collected at the same location as the holotype on 13 June 2012; one adult male (AMS R186113), collected near the type locality, beside a 2 m wide swiftly cascading stream (22.31387�� N, 103.7654�� E, 2793 m asl; Fig 1), collected on 11 June 2012. All aforementioned specimens collected by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. A gravid female (AMS R186116) and two calling adult males (AMS R186114 & AMS R186115), collected from disturbed bamboo and elfin forest near a steep rocky stream on Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam, (22.31412�� N, 103.76588�� E, 2764 m asl; Figs. 1 & 6C), collected between 20:15 and 20:20 h on 21 June 2016 by Jodi J. L. Rowley, Benjamin Tapley and Nguyen Thanh Chung. Referred specimen: One adult male (HLNP 20120620 00002), found resting on leaf litter within a bamboo forest beside a 1 m wide cascading stream on Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam (22.31349�� N, 103.76521�� E, 2813 m asl; Fig. 1), collected on 10 June 2012 by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. Etymology: Specific epithet ��� fansipanensis ��� is a toponym in reference to the type locality of the species, meaning ���from [Mount] Fansipan���. Suggested vernacular name: Mount Fansipan horned frog (English), ���ch s���ng phan xi păng (Vietnamese). Diagnosis: Megophrys fansipanensis sp. nov. differs from its congeners by a combination of the following morphological characters: (1) small size, adult male SVL 30.9���44.3 mm (N =13), adult female SVL 41.7���42.5 mm (N =2); (2) small palpebral horn on upper eyelid; (3) toes lacking distinct interdigital webbing and lateral dermal fringes indistinct or absent; (4) TYD/EL 53.0���80.0% in adult males (N =13), 63.0���71.0% in adult females (N =2); (5) subarticular tubercles absent; (6) palmar tubercles absent; (7) inner metatarsal tubercle present; (8) SHL/SVL 49.0���58.9% in adult males (N =13), 55.8���58.5% in adult females (N =2); (9) tympanum clearly defined; (10) vomerine ridges, vomerine and maxillary teeth present; and (11) an advertisement call with an average dominant frequency of 3.8 kHz at 15.3���18.3 ��C. Description of holotype (measurements provided in Table 2): Sexually mature male (Figs. 3 A���B, 4A���C, 5A & 5D). Head small, longer than wide; snout rounded in dorsal view, obtusely protruding in lateral view, rostral appendage absent; loreal region vertical and concave; canthus rostralis angular; dorsal region of the snout slightly concave; eye length 33% greater than the maximum diameter of the tympanum and subequal to the snout length; eye to tympanum distance shorter than the maximum tympanum diameter; tympanum circular. Pupil in life oval, vertically orientated when dilated; nostrils orientated laterally, equidistant to the eyes and the snout; internarial distance is less than the width of the upper eyelids, and less than the narrowest point between the upper eyelids; upper eyelid width and the shortest distance between the upper eyelids are subequal to one another. Vomerine ridges present, oblique and barely separated anteriorly, with small vomerine teeth; maxillary teeth present. Tongue moderately large and not clearly notched posteriorly. Forelimbs moderately long and thin, forearm length shorter than the hand length; fingers long and narrow, without lateral fringes, finger length formula FIColour of holotype: In life (Fig. 5A & 5D): Dorsally dark tan; darker lines follow the opposing ���V���-shaped parietoscapular-sacral ridges; dark brown loreal region, snout pale; darker brown triangular marking between the eyes with a lighter central blotch; vertical dark brown bar below the eyes; dark brown tympanum; dark tan hind and forelimbs possess dark bars; supratympanic fold longitudinally bicolored, cream above and dark brown below; tubercles in the inguinal region either a pale cream or dark brown in colour; gular and pectoral region dark greybrown, gular region with a medial dark brown stripe; abdomen lighter with dark blotches and speckling; ventral surface of the thighs speckled with brown; iris metallic orange with black reticulations throughout. In preservative (Figs. 3 A���B & 4A���C): The majority of the dorsal and lateral surfaces of the head, body, forelimbs and hindlimbs brown; darker brown triangular marking with light central blotch between the eyes; darker brown ���X���-shaped marking over the opposing ���V���-shaped parietoscapular-sacral ridges; dorsolateral and supratympanic folds and flank tubercles brownish-cream; the front of the snout and the lateral canthus rostralis dark brown; wide vertical dark brown bar below the eyes; dark brown tympanum; two dark brown blotches on the anterior lateral surface of the forearms; dorsal surface of FII, FIII, FIV with dark brown blotches. The gular region, chest and the anterior part of the abdomen primarily creamy-grey, with brown speckling on the gular region; gular region with a medial dark brown stripe. Abdomen blotched with dark brown, large brown blotches incompletely encircle the abdomen; ventral surfaces of the thighs and shanks with pale brown mottling; ventral surfaces of the feet grey-brown; area surrounding the vent dark brown; forelimbs ventrally mottled and blotched with light and dark brown; ventral surface of the hands grey; pectoral and femoral glands white. Refer to Figures 3 A���B, 4A���C, 5A & 5D for further details of markings and colouration. Variation: Morphometric measurements of the type series and referred specimen are given in Table 2. Paratypes and the referred specimen generally agree with the holotype morphologically, but with the following exceptions: Head width greater than the head length for 11 of the 15 specimens (VNMN 0 7033, AMS R186106��10, AMS R186112, AMS R186113, AMS R186115, AMS R186116 & HLNP 20120620 00002); head width less than the head length for two specimens (VNMN 2018.01 & AMS R186105), and head length equal to the head width for two specimens (AMS R186111 & AMS R186114). Seven of the 15 specimens (VNMN 2018.01, AMS R186106 & AMS R186109 ��13) lack dermal fringes on the toes whereas indistinct ridges were observed on the toes of five specimens (VNMN 0 7033, AMS R186105, AMS R186107, AMS R186108 & HLNP 20120620 00002). It is not clear if the presence of indistinct ridges on the toes of these specimens is an artefact of specimen positioning during the fixation process. Relative finger lengths differ from the holotype as follows: FISecondary sexual characters: All males had slightly raised nuptial pads covered with black microspinules; nuptial pads covering most of the dorsal surface of the base of FI and FII; nuptial pad oval shaped on FI and FII; a protruding fleshy projection posterior to the cloaca (a secondary sexual character of some male Megophrys, e.g., M. caudoprocta Shen, 1994, M. koui and M. pachyproctus) is absent on all male specimens. Female (AMS R186116) with large unpigmented ova; both examined females did not possess nuptial pads or dermal asperities. Molecular data: 16S sequences were generated from three adult specimens. Uncorrected p -distance between Megophrys fansipanensis sp. nov. and other Megophrys sequences available on GenBank were ��3.3%. Advertisement call: Call descriptions are based on the call groups of the holotype (VNMN 2018.01, N= 3) and two paratypes (AMS R186114 [N= 12] & AMS R186115 [N= 5]), and the total number of calls analysed (N =60) was 20 calls per individual. Advertisement calls were recorded at 15.3���18.3 ��C ambient temperature. Calls were an average of 42 ms (34���49 ms) in duration (Table 3, Fig. 7). Calls contained an average of 11.6 pulses (10���14). The average dominant frequency of all analysed calls was 3.8 kHz (3.6���4.7 kHz). Calls were repeated at a rate of 3.9 (3.8���4.0) calls per second in the analysed call groups and had an average intercall interval of 204.4 ms (180���290 ms). The number of calls within each call group ranged from 1 to 41, with an average of 22.8. Most call groups began with one to three calls at relatively low amplitude, with amplitude increasing with each call, after which amplitude remained relatively constant (Fig. 7). Individual calls decreased in amplitude over time (Fig. 7). Natural history: Megophrys fansipanensis sp. nov. is associated with upper montane bamboo forest, mixed bamboo forest and disturbed secondary broadleaf forest with a relatively open canopy. All individuals have been encountered at night and found in riparian habitat along 1���2 m wide streams with clear water and rocky stream beds. The majority of individuals were encountered in the month of June, fewer animals were observed in September. Males were calling from stream-side vegetation in June 2012 and June 2016. A gravid female was found adjacent to a fast-flowing stream in June 2016. Tadpoles were not observed at this time. During field work in September 2015, only a single unvouchered male was encountered, no vocalisations were heard and tadpoles were not observed. AMS R186115 VNMN 2018.01 AMS R186114 Number of call groups measured 5 3 12 Number of calls measured 20 20 20 Call duration (ms) 41.2 (38���45) 43.8 (41���49) 40.8 (34���45) Intercall interval (ms) 213.7 (197���245) 197.7 (180���227) 201.8 (168���290) Call repetition rate (calls/s) 3.8 4.0 4.0 Calls/call group 22.2 (18���25) 34.7 (24���41) 11.4 (1���22) Pulses/call 11.1 (10���12) 10.8 (10���11) 12.9 (12���14) Dominant frequency (kHz) 3.6 (na) 3.9 (3.9���4.1) 4.0 (3.6���4.7) Temperature (��C) 15.3 18.3 15.3 Distribution and conservation status: Megophrys fansipanensis sp. nov. is only known from Mount Fansipan at elevations between 2200 and 2813 m asl (Fig. 1). We did not detect this species during field work on Mount Fansipan at elevations of 1200 m asl (June 2016), 1700 m asl (June 2012 & 2016) and 1900 m asl (June 2016; September 2017). The high elevation sites where this species occurs face the immediate threat of habitat degradation and pollution due to the impact of tourists; habitats were noticeably polluted with garbage and runoff from toilets near the stream (Fig. 6D). In addition, if Megophrys fansipanensis sp. nov. is restricted to a narrow, high-elevation band, then the species is likely to be particularly vulnerable to climate change. Megophrys fansipanensis sp. nov. may occur more widely, but it was not detected in habitat similar to that found on Mount Fansipan during recent surveys on Mount Ky Quan San, 25 km north of our collection sites. The current EOO is predicted to be 630 km 2, qualifying Megophrys fansipanensis nov. sp. for listing as Endangered in accordance with the IUCN Red List of Threatened Species categories and criteria B1ab(iii). Comparisons: Megophrys fansipanensis sp. nov. can be distinguished from all species in the subgenus Panophrys found in mainland Southeast Asia, north of the Isthmus of Kra and neighbouring provinces of China on the basis of morphology, and from all species in the subgenus for which comparable data is available on the basis of molecular and acoustic data. Comparisons with each subgenus are discussed separately below. The following comparison is based on 13 males and two females of Megophrys fansipanensis sp. nov. Subgenus Brachytarsophrys: Megophrys fansipanensis sp. nov. can be distinguished from M. carinense, M. feae and M. intermedia by the absence of a transverse fold at the base of the head (versus presence in M. carinense, M. feae and M. intermedia) and having a smaller adult male size, SVL 30.9���44.3 mm (versus SVL> 79.1 mm; material examined). Subgenus Ophryophryne: Megophrys fansipanensis sp. nov. can be distinguished from all species in the subgenus Ophryophryne due to the presence (versus absence) of maxillary teeth (Poyarkov et al. 2017; material examined) and further from M. elfina by having a larger adult female size, SVL 41.7���42.5 mm (versus 35.1���36.5 mm, N =6, in M. elfina; Poyarkov et al. 2017) and the presence of vomerine teeth (versus absence in M. elfina; Poyarkov et al. 2017); from M. gerti by having a smaller adult female size, SVL 41.7���42.5 mm (versus 43.1���47.4 mm, N =3, in M. gerti; Poyarkov et al. 2017; material examined) and the presence of vomerine ridges and teeth (versus absence in M. gerti; Poyarkov et al. 2017); from M. hansi by having a smaller adult female size, SVL 41.7��� 42.5 mm (versus 45.1���53.9 mm, N =5, in M. hansi; Poyarkov et al. 2017; material examined); from M. koui by the absence of a protruding fleshy projection above the cloaca in sexually mature males (secondary sexual characteristic of sexually mature M. koui; Poyarkov et al. 2017); and from M. synoria by having a smaller adult female size, SVL 41.7���42.5 mm (versus 51.4���70.7 mm, N =3, in M. synoria; Poyarkov et al. 2017) and the presence of microspinules on nuptial pads of sexually mature males (versus microgranules on nuptial pads of sexually mature male M. synoria; material examined). Subgenus Xenophrys: Megophrys fansipanensis sp. nov. differs from M. auralensis by having a smaller adult male size, SVL 30.9���44.3 mm (versus 60.1�� 76.9 mm, N =20, in M. auralensis; Ohler et al. 2002; Neang et al. 2013); from M. damrei by having a smaller adult male size, SVL 30.9���44.3 mm (versus 47.7���57.1 mm, N =7, in M. damrei; Mahony 2011; Neang et al. 2013; material examined); from M. glandulosa by having a smaller adult male size, SVL 30.9���44.3 mm (versus 76.7���81.6 mm, N =10, in M. glandulosa; Fei et al. 2009; material examined), a lack of dermal fringes on toes (versus presence in M. glandulosa; Huang et al. 1998; Mahony et al. in press), a lack of distinct interdigital webbing between toes (versus basal webbing in M. glandulosa; material examined), and the absence of a light-coloured upper lip stripe (versus presence in M. glandulosa; Fei et al. 1990; Mahony et al. in press); from M. lekaguli by having a smaller adult male size, SVL 30.9���44.3 mm (versus 55.6���68.1 mm, N =8, in M. lekaguli; Stuart et al. 2006b; material examined); from M. major by having a smaller adult male size, SVL 30.9&nd, Published as part of Tapley, Benjamin, Cutajar, Timothy, Mahony, Stephen, Nguyen, Chung Thanh, Dau, Vinh Quang, Luong, Anh Mai, Le, Dzung Trung, Nguyen, Tao Thien, Nguyen, Truong Quang, Portway, Christopher, Luong, Hao Van & Rowley, Jodi J. 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Megophrys (Panophrys) hoanglienensis sp. nov. Figs. 8–13 & Table 4. Holotype: One adult male (VNMN 2018.02), found beside a 5 m wide low-gradient shallow stream in slightly disturbed evergreen forest, Mount Fansipan, Hoang Lien National Park (HLNP), Sa Pa District, Lao Cai Province, Vietnam (22.33518° N, 103.77947° E, 2075 m asl; Fig. 8), collected at 19:15 h on 14 June 2012 by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. Paratypes: Four adult males (VNMN 0 7034, AMS R186118, AMS R186119 & AMS R186120), found in the vicinity of a 2 m wide low-gradient stream in disturbed evergreen forest, Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam (22.32827° N, 103.78184° E, 2242 m asl; Fig. 8), collected at 19:45 h on 12 June 2012; one adult female (AMS R186121), found beside a 5 m wide low-gradient shallow stream in slightly disturbed evergreen forest, Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam (22.33518° N, 103.77947° E, 2075 m asl), collected at 19:15 h on 14 June 2012. All aforementioned specimens collected by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. Two adult males (AMS R186122 & AMS R186123), found calling from a rocky overhang 1.8 m above and 1.5 m away from a 3 m wide cascading stream at the base of Love Waterfall (thác Tình Yêu) in disturbed evergreen forest on Mount Fansipan, HLNP, Tam Duong District, Lai Chau Province, Vietnam (22.35065° N, 103.77136° E, 1898 m asl; Figs. 8 & 12 A–B), collected at 22:15 h on 24 June 2016 by Jodi J. L. Rowley, Benjamin Tapley, Cong Huy Le and Nguyen Thanh Chung; three adult males (AMS R186125, AMS R186126 & AMS R186127), found beside a swampy stream in disturbed evergreen forest, Mount Ky Quan San, Bat Xat District, Lao Cai Province, Vietnam (22.30343° N, 103.37019° E, 2153 m asl), collected at 21:00 h on 0 6 September 2017 by Nguyen Thanh Chung, Luan Thanh Nguyen, Luke Harding, Timothy Cutajar, Jodi J. L. Rowley and Benjamin Tapley. Referred specimens: One adult male (HLNP 20120624 00003), found calling from a rocky overhang 1.8 m above and 1.5 m away from a 3 m wide cascading stream at the base of Love Waterfall in disturbed evergreen forest on Mount Fansipan, HLNP, Tam Duong District, Lai Chau Province, Vietnam (22.35065° N, 103.77136° E, 1898 m asl; Figs. 8 & 12 A–B), collected at 22:15 h on 24 June 2016 by Jodi J. L. Rowley, Benjamin Tapley, Cong Huy Le and Nguyen Thanh Chung; two juvenile specimens (HLNP 20170906 0 0 0 0 4 & AMS R186124), found beside a 3 m wide stream in disturbed evergreen forest on Mount Fansipan, HLNP, Tam Duong District, Lai Chau Province, Vietnam (22.21039° N, 103.46293° E, 1882 m asl), collected at 20:00 h on 0 6 September 2017 by Nguyen Thanh Chung, Luan Thanh Nguyen, Luke Harding, Timothy Cutajar, Jodi J. L. Rowley and Benjamin Tapley; six adult males (HNUE MCC.2017.83–85, HNUE MCC.2017.88 & IEBR MCC.2017.86–87), found in the vicinity of Hang De Cha Hai stream, Che Tao Village, Che Tao Commune, Mu Cang Chai District, Mu Cang Chai Species Habitat Conservation Area, Yen Bai Province, Vietnam (21.76247° N, 104.01889° E, 2046 m asl), collected on 0 1 May 2017 by Yen Thi Do, Anh Ngoc Dao and Nam Hai Nguyen. Etymology: Specific epithet “ hoanglienensis ” is a toponym in reference to the type locality of the species, meaning “from Hoang Lien” Range. Suggested vernacular name: Hoang Lien horned frog (English), Ếch sừng hoàng liên (Vietnamese). Diagnosis: Megophrys hoanglienensis sp. nov. differs from its congeners by a combination of the following morphological characters: (1) medium adult size, adult male SVL 37.4–47.6 mm (N= 11); adult female SVL 59.6 mm (N= 1); (2) small palpebral horn on upper eyelid present; (3) interdigital webbing and lateral fringes on the toes indistinct or absent; (4) snout mucronate in dorsal view; (5) TYD/EL 54.2–75.0% in adult males (N =11), 60.0% in adult female (N= 1); (6) subarticular tubercles absent; (7) palmar tubercles absent; (8) inner metatarsal tubercle present; (9) HW/SVL 37.4–42.5% in adult males (N= 11), 38.3% in adult female (N= 1); (10) SHL/SVL 44.3– 62.5% in adult males (N =11), 57.5% in adult female (N= 1); (11) tympanum clearly defined; (12) TYE/SVL 6.3– 9.5% in adult males (N= 11), 7.6% in adult female (N= 1); (13) vomerine ridges, vomerine and maxillary teeth present; (14) an advertisement call with a dominant frequency of 2.8–3.0 kHz at 18.5 °C. Description of holotype (measurements provided in Table 4): Sexually mature male (Figs. 9 A–B, 10A–C, 11A–B). Head small, wider than long; snout mucronate in dorsal view, obtusely protruding in lateral view, rostral appendage absent; loreal region vertical and concave; canthus rostralis angular; dorsal region of the snout slightly concave; eye length 38.0% longer than the maximum tympanum diameter and subequal to the snout length; eye to tympanum distance greater than the maximum tympanum diameter; tympanum circular. Pupil in life oval, vertically orientated when dilated; nostrils orientated laterally, closer to the eyes than the snout; internarial distance subequal to the width of the eyelids, and subequal to the narrowest point between the upper eyelids; vomerine ridges present, oblique and barely separated anteriorly, with small vomerine teeth; maxillary teeth present. Tongue moderately large and feebly notched posteriorly. Forelimbs moderately long and thin; forearm length shorter than the hand length; fingers long and narrow without lateral fringes, finger length formula FI=FII=FIV F1 GURE 10. Megophrys hoanglienensis sp. nov. in preservative (holotype VNMN 2018.02). (A) Lateral view of head, (B) palmar surface of right hand, and (C) plantar surface of left foot. 5 mm scale bar. Skin of dorsal surfaces of the body, limbs, and both dorsal and lateral surfaces of the head weakly granular; tympanum granular with borders slightly raised; very small palpebral horn present on the outer edge of the upper eyelid; supratympanic fold narrows as it passes above the tympanum, widening again before terminating above the axilla; flanks with small scattered tubercles; thin dorsolateral fold extending posteriorly from behind the supratympanic fold to the groin on each side; a weak, “V”-shaped parietoscapular ridge present, its two sides extending posteriorly from above the tympanum and meeting medially beyond the level of the axilla; a second weak inverted “U”-shaped sacral ridge present on the mid-dorsum which joins laterally with the dorsolateral folds; “V” and “U”-shaped ridges not joined at their apex, apex of ridges with black asperities; small tubercles present on the dorsal surface of the thighs, shanks and forearms. Large distinct tubercles present on the flanks, inguinal region and dorsal surface of the hindlimbs; chest and ventral surfaces of the forelimbs and shanks smooth; pectoral glands obvious, small, slightly raised, positioned level with the axilla; femoral glands small, slightly raised, one positioned equidistant from the knee and the cloaca on the posterior surface of each thigh; white skin asperities forming narrow band circummarginally on the upper and lower jaw; black asperities on the posterior parts of the upper eyelids, on the dorsum and area directly above the vent, and on tubercles of the tympanic regions, absent elsewhere. Colour of holotype: In life (Figs. 11 A–B): Dorsally dark tan; with light tan blotches; darker brown triangular marking between the eyes with a lighter central blotch; cream supratympanic folds with black lower margin; inguinal regions light tan; dark brown hindlimbs with darker blotches; posterior surface of the thighs and shanks very dark brown; forearms light brown with a single dark bar; dorsal surface of the fingers with dark brown bars; gular, pectoral and abdomen region mottled grey-brown and cream, gular region with a medial dark brown stripe; ventral surface of thighs unmarked; pectoral glands off-yellow; iris metallic gold. In preservative (Figs. 9 A–B & 10A–C): Markings are the same as in life, colouration differences are as follows: The majority of the dorsal and lateral surfaces of the head, body, forelimbs and hindlimbs brown; dorsolateral and supratympanic folds and flank tubercles brownish-cream; tip of the snout cream, lateral canthus rostralis dark brown; wide vertical dark brown bar below the eyes; dark brown band running from the posterior edge of the eyes, through the tympanum, and terminating where the forelimbs join the body; lower margin of the supratympanic folds dark brown, sharply contrasting with creamy grey upper margin; gular region, chest and anterior part of the abdomen primarily creamy-grey with brown speckling, and a dark brown medial bar in the gular region; dark brown blotches anterior to each of the pectoral glands, pectoral glands light cream in colour; abdomen speckled with dark brown; ventral surfaces of the thighs and shanks with diffuse pale brown mottling; ventral surfaces of the feet grey brown with cream digit tips; area surrounding the vent dark brown; forelimbs ventrally mottled with dark brown bar running longitudinally on the outer ventral surface of the forearms; grey colouration of ventral side of the hands with darker mottling and cream digit tips; femoral glands cream. Variation: Morphometric measurements of the type series and referred specimens are provided in Table 4. Nuptial pads were not visible on one male specimen (AMS VNMN 07034) although sex was confirmed via inspection of the gonads. Two (AMS R186120 & AMS R186121) of the 12 specimens possessed rudimentary webbing between the toes. Four (VNMN 0 7034 & AMS R186118¯20) of the 12 specimens with indistinct lateral fringes on the toes. It is not clear if the presence of indistinct fringes on the toes of these specimens is an artefact of the positioning during the fixation process. Relative finger lengths are highly variable, and only one specimen (AMS R186119; FIẼFII Secondary sexual characters: All sexually mature males had slightly raised nuptial pads covered with black microspinules, nuptial pads covering most of the dorsal surface of the base of FI and FII; nuptial pad oval shaped on FI and FII; the protruding fleshy projection posterior to the cloaca (a secondary sexual character of some male Megophrys, e.g., M. caudoprocta, M. koui and M. pachyproctus) is absent on all male specimens. The examined female did not possess nuptial pads. Molecular data: Sequences generated from nine adult specimens in this study (Table 1) were most similar to “ Megophrys sp. 5” (sensu Chen et al. 2017) collected in Sa Pa District, Lao Cai Province, Vietnam; 16S rDNA sequences differed by only 0.3% uncorrected p -distance between “ Megophrys sp. 5” and our specimens (from Lao Cai & Lai Chau provinces), and by 1.1% for specimens collected approximately 58 km away in Yen Bai Province, which is greater than the intraspecific variation reported elsewhere within the subgenus Panophrys (e.g., Li et al. 2014 reported an intraspecific p -distance of 0.6% between populations of M. obesa Wang, Li & Zhao, 2014), but is lower than intraspecific variation within Megophrys sensu lato (e.g., 3.1% mean uncorrected p -distance in M. elfina; Poyarkov et al. 2017). Uncorrected p -distance was 0.8% between specimens of Megophrys hoanglienensis sp. nov. collected from Lao Cai and Lai Chau provinces. The smallest interspecific p- distance in our study was between Megophrys hoanglienensis sp. nov. and Megophrys fansipanensis sp. nov. at 3.3% where the two new species occur in syntopy supporting their position as distinct sister taxa. Advertisement call: Call descriptions are based on the calls of one paratype (AMS R186122), five call groups and 20 calls in total were analysed. Advertisement calls were recorded at 18.5 °C ambient temperature. Calls were an average of 102.9 ms (96–108 ms) in duration (Table 5; Fig. 13). The average number of pulses per call was 18.7 (12–22). The average dominant frequency of all the analysed calls was 3.0 kHz (2.8–3.0 kHz). Calls were repeated at a rate of 2.6 calls per second for the call groups analysed, and had an average intercall interval of 274.2 ms (178– 565 ms). The number of calls within each call group ranged from 11 to 21, with an average of 17. Most call groups began at a relatively low amplitude, increasing with each call up to approximately a quarter of the duration, after which amplitude varied but did not follow a trend or pattern (Fig. 13A & B). Individual calls began with a medium relative amplitude but peaked early before declining towards the end, and the interval between the first and second pulse was longer than those between any of the following pulses (Fig. 13C). Natural history: Megophrys hoanglienensis sp. nov. is associated with disturbed secondary broadleaf forest with a relatively open canopy. All individuals have been found at night in riparian habitat along clear water, 2–5 m wide streams with a gravel or rocky substrate. Males were calling from streamside vegetation in June 2012 and June 2016 but not during the surveys undertaken in May or September 2017. A non-gravid female was found adjacent to a wide, low-gradient, shallow stony stream in June 2016. Tadpoles were not observed at any of the locations where this species has been found (surveys undertaken in June 2012 and 2016, and in September 2015 and 2017). Megophrys hoanglienensis sp. nov. is sympatric with M. jingdongensis at 1990 m asl on Mount Fansipan, and with M. fansipanensis sp. nov. on Mount Fansipan at 2242 m asl. Megophrys hoanglienensis sp. nov. is sympatric with M. jingdongensis and M. rubrimera on Mount Ky Quan San at 2153 m asl (Tapley et al. 2018b). Distribution and conservation status: This species is known from between 1898–2242 m asl at four localities, up to 92 km apart in the Hoang Lien Range (Sa Pa District, Lao Cai Province, HLNP; Tam Duong District, Lai Chau Province; Mount Ky Quan San, Bat Xat District, Lao Cai Province and Che Tao Commune, Yen Bai Province, Vietnam). The species’ EOO is currently predicted to be 3216 km 2 (Fig. 8). Ongoing disturbance to the species’ habitat due to forest clearance for agriculture has been observed in the parts of Sa Pa District where the species was observed. In Bat Xat District the forest in which this species occurs is being negatively impacted by fuelwood collection for the tourism industry and by the grazing of livestock. Megophrys hoanglienensis sp. nov. is likely to qualify as Endangered in accordance with the IUCN Red List of Threatened Species categories and criteria B1ab(iii). Comparisons: Megophrys hoanglienensis sp. nov. can be distinguished from all species in the subgenus Panophrys found in mainland Southeast Asia, north of the Isthmus of Kra and neighbouring provinces of China on the basis of morphology, and from all species in the subgenus for which comparable data is available on the basis of molecular and acoustic data. Comparisons with each subgenus are discussed separately below. The following comparison is based on 11 males and one female of Megophrys hoanglienensis sp. nov. Subgenus Brachytarsophrys: Megophrys hoanglienensis sp. nov. can be distinguished from M. carinense, M. feae and M. intermedia by the absence of a transverse fold at the base of the head (versus presence in M. carinense, M. feae and M. intermedia), and by having a smaller adult male size, SVL 41.1–47.6 mm (versus SVL> 79.1 mm; material examined). Subgenus Ophryophryne: Megophrys hoanglienensis sp. nov. can be distinguished from all species in the subgenus Ophryophryne due to the presence (versus absence) of maxillary teeth (Poyarkov et al. 2017; material examined); further from M. elfina by having a larger adult female size, SVL 59.6 mm (versus 35.1–36.5 mm, N =6, in M. elfina; Poyarkov et al. 2017); from M. gerti by having a larger adult female size, SVL 59.6 mm (versus 43.1– 47.4 mm, N =3, in M. gerti; Poyarkov et al. 2017); from M. hansi by having a larger adult female size, SVL 59.6 mm (versus 45.1–53.9 mm, N =5, in M. hansi; Poyarkov et al. 2017; material examined); from M. koui by the absence of a protruding fleshy projection above the cloaca in sexually mature males (secondary sexual characteristic of adult M. koui; Poyarkov et al. 2017); and from M. synoria by the presence of microspinules on nuptial pads in adult males (versus microgranules on nuptial pads in adult M. synoria; Poyarkov et al. 2017; material examined). Subgenus Xenophrys: Megophrys hoanglienensis sp. nov. differs from M. auralensis by having a smaller adult male body size, SVL 37.4–47.6 mm (versus 60.1–76.9 mm, N = 20 in M. auralensis, N =20; Ohler et al. 2002; Neang et al. 2013); from M. damrei by having a smaller adult female body size, SVL 59.6 mm (versus 69.1 mm, N= 1, in M. damrei; Mahony 2011; material examined); from M. glandulosa by having a smaller adult male body size, SVL 37.4–47.6 mm (versus 76.7–81.6 mm, N =10, in M. glandulosa; Fei et al. 2009; Fei et al. 1990; material examined), absence of distinct lateral fringes on toes (versus presence in M.

Megophrys (Panophrys) fansipanensis sp. nov. Figs. 1, 3–7 & Table 2. Holotype: Adult male (VNMN 2018.01), found perched on a stick 3 m above a 2 m wide low-gradient stream in disturbed evergreen forest on Mount Fansipan, Hoang Lien National Park (HLNP), Sa Pa District, Lao Cai Province, Vietnam (22.32827° N, 103.78184° E, 2242 m asl; Fig. 1), collected at 19:15 h on 12 June 2012 by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. Paratypes: Adult female (VNMN 07033), found resting on leaf litter associated with bamboo forest beside a 1 m wide cascading stream on Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam (22.31458° N, 103.76574° E, 2783 m asl; Fig 1), collected at 20:00 h on 0 9 June 2012; four adult males (AMS R186105–08), found in the vicinity of a 1 m wide cascading stream on Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam (22.31349° N, 103.76521° E, 2813 m asl; Fig. 1), on 10 June 2012; four adult males (AMS R186109–12), collected at the same location as the holotype on 13 June 2012; one adult male (AMS R186113), collected near the type locality, beside a 2 m wide swiftly cascading stream (22.31387° N, 103.7654° E, 2793 m asl; Fig 1), collected on 11 June 2012. All aforementioned specimens collected by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. A gravid female (AMS R186116) and two calling adult males (AMS R186114 & AMS R186115), collected from disturbed bamboo and elfin forest near a steep rocky stream on Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam, (22.31412° N, 103.76588° E, 2764 m asl; Figs. 1 & 6C), collected between 20:15 and 20:20 h on 21 June 2016 by Jodi J. L. Rowley, Benjamin Tapley and Nguyen Thanh Chung. Referred specimen: One adult male (HLNP 20120620 00002), found resting on leaf litter within a bamboo forest beside a 1 m wide cascading stream on Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam (22.31349° N, 103.76521° E, 2813 m asl; Fig. 1), collected on 10 June 2012 by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. Etymology: Specific epithet “ fansipanensis ” is a toponym in reference to the type locality of the species, meaning “from [Mount] Fansipan”. Suggested vernacular name: Mount Fansipan horned frog (English), Ếch sừng phan xi păng (Vietnamese). Diagnosis: Megophrys fansipanensis sp. nov. differs from its congeners by a combination of the following morphological characters: (1) small size, adult male SVL 30.9–44.3 mm (N =13), adult female SVL 41.7–42.5 mm (N =2); (2) small palpebral horn on upper eyelid; (3) toes lacking distinct interdigital webbing and lateral dermal fringes indistinct or absent; (4) TYD/EL 53.0–80.0% in adult males (N =13), 63.0–71.0% in adult females (N =2); (5) subarticular tubercles absent; (6) palmar tubercles absent; (7) inner metatarsal tubercle present; (8) SHL/SVL 49.0–58.9% in adult males (N =13), 55.8–58.5% in adult females (N =2); (9) tympanum clearly defined; (10) vomerine ridges, vomerine and maxillary teeth present; and (11) an advertisement call with an average dominant frequency of 3.8 kHz at 15.3–18.3 °C. Description of holotype (measurements provided in Table 2): Sexually mature male (Figs. 3 A–B, 4A–C, 5A & 5D). Head small, longer than wide; snout rounded in dorsal view, obtusely protruding in lateral view, rostral appendage absent; loreal region vertical and concave; canthus rostralis angular; dorsal region of the snout slightly concave; eye length 33% greater than the maximum diameter of the tympanum and subequal to the snout length; eye to tympanum distance shorter than the maximum tympanum diameter; tympanum circular. Pupil in life oval, vertically orientated when dilated; nostrils orientated laterally, equidistant to the eyes and the snout; internarial distance is less than the width of the upper eyelids, and less than the narrowest point between the upper eyelids; upper eyelid width and the shortest distance between the upper eyelids are subequal to one another. Vomerine ridges present, oblique and barely separated anteriorly, with small vomerine teeth; maxillary teeth present. Tongue moderately large and not clearly notched posteriorly. Forelimbs moderately long and thin, forearm length shorter than the hand length; fingers long and narrow, without lateral fringes, finger length formula FI Skin of the dorsal surfaces of the body, limbs and dorsal and lateral surfaces of the head weakly granular; tympanum granular with borders slightly raised; very small palpebral horn present on the outer edge of the upper eyelids; supratympanic fold narrows as it passes above the tympanum and expands in width again posteriorly, terminating above the axilla; flanks with small scattered tubercles; thin dorsolateral fold on each side, extending from behind the supratympanic fold to approximately two-thirds of the distance to the groin; a weak, “V”-shaped parietoscapular ridge present, its two sides extending posteriorly from above the tympanum and meeting medially beyond the level of the axilla; a second inverted “V”-shaped sacral ridge present on the mid-dorsum which joins laterally with the dorsolateral folds; “V”-shaped ridges joined at their apices by a medial ridge; small tubercles arranged into distinct transverse rows on the dorsal surfaces of thighs, shanks and forearms. Large distinct tubercles present on the inguinal region; gular region, chest and ventral surfaces of the limbs smooth; pectoral glands distinct, small, slightly raised, positioned level with the axilla; femoral glands small, slightly raised, one positioned equidistant from the knee and cloaca on the posterior surface of each thigh; white skin asperities form a narrow band circummarginally on the upper and lower jaw; black asperities on the tubercles of the tympanic regions and posterior parts of the upper eyelids, absent elsewhere. Colour of holotype: In life (Fig. 5A & 5D): Dorsally dark tan; darker lines follow the opposing “V”-shaped parietoscapular-sacral ridges; dark brown loreal region, snout pale; darker brown triangular marking between the eyes with a lighter central blotch; vertical dark brown bar below the eyes; dark brown tympanum; dark tan hind and forelimbs possess dark bars; supratympanic fold longitudinally bicolored, cream above and dark brown below; tubercles in the inguinal region either a pale cream or dark brown in colour; gular and pectoral region dark greybrown, gular region with a medial dark brown stripe; abdomen lighter with dark blotches and speckling; ventral surface of the thighs speckled with brown; iris metallic orange with black reticulations throughout. In preservative (Figs. 3 A–B & 4A–C): The majority of the dorsal and lateral surfaces of the head, body, forelimbs and hindlimbs brown; darker brown triangular marking with light central blotch between the eyes; darker brown “X”-shaped marking over the opposing “V”-shaped parietoscapular-sacral ridges; dorsolateral and supratympanic folds and flank tubercles brownish-cream; the front of the snout and the lateral canthus rostralis dark brown; wide vertical dark brown bar below the eyes; dark brown tympanum; two dark brown blotches on the anterior lateral surface of the forearms; dorsal surface of FII, FIII, FIV with dark brown blotches. The gular region, chest and the anterior part of the abdomen primarily creamy-grey, with brown speckling on the gular region; gular region with a medial dark brown stripe. Abdomen blotched with dark brown, large brown blotches incompletely encircle the abdomen; ventral surfaces of the thighs and shanks with pale brown mottling; ventral surfaces of the feet grey-brown; area surrounding the vent dark brown; forelimbs ventrally mottled and blotched with light and dark brown; ventral surface of the hands grey; pectoral and femoral glands white. Refer to Figures 3 A–B, 4A–C, 5A & 5D for further details of markings and colouration. Variation: Morphometric measurements of the type series and referred specimen are given in Table 2. Paratypes and the referred specimen generally agree with the holotype morphologically, but with the following exceptions: Head width greater than the head length for 11 of the 15 specimens (VNMN 0 7033, AMS R186106¯10, AMS R186112, AMS R186113, AMS R186115, AMS R186116 & HLNP 20120620 00002); head width less than the head length for two specimens (VNMN 2018.01 & AMS R186105), and head length equal to the head width for two specimens (AMS R186111 & AMS R186114). Seven of the 15 specimens (VNMN 2018.01, AMS R186106 & AMS R186109 ¯13) lack dermal fringes on the toes whereas indistinct ridges were observed on the toes of five specimens (VNMN 0 7033, AMS R186105, AMS R186107, AMS R186108 & HLNP 20120620 00002). It is not clear if the presence of indistinct ridges on the toes of these specimens is an artefact of specimen positioning during the fixation process. Relative finger lengths differ from the holotype as follows: FI Secondary sexual characters: All males had slightly raised nuptial pads covered with black microspinules; nuptial pads covering most of the dorsal surface of the base of FI and FII; nuptial pad oval shaped on FI and FII; a protruding fleshy projection posterior to the cloaca (a secondary sexual character of some male Megophrys, e.g., M. caudoprocta Shen, 1994, M. koui and M. pachyproctus) is absent on all male specimens. Female (AMS R186116) with large unpigmented ova; both examined females did not possess nuptial pads or dermal asperities. Molecular data: 16S sequences were generated from three adult specimens. Uncorrected p -distance between Megophrys fansipanensis sp. nov. and other Megophrys sequences available on GenBank were ±3.3%. Advertisement call: Call descriptions are based on the call groups of the holotype (VNMN 2018.01, N= 3) and two paratypes (AMS R186114 [N= 12] & AMS R186115 [N= 5]), and the total number of calls analysed (N =60) was 20 calls per individual. Advertisement calls were recorded at 15.3–18.3 °C ambient temperature. Calls were an average of 42 ms (34–49 ms) in duration (Table 3, Fig. 7). Calls contained an average of 11.6 pulses (10–14). The average dominant frequency of all analysed calls was 3.8 kHz (3.6–4.7 kHz). Calls were repeated at a rate of 3.9 (3.8–4.0) calls per second in the analysed call groups and had an average intercall interval of 204.4 ms (180–290 ms). The number of calls within each call group ranged from 1 to 41, with an average of 22.8. Most call groups began with one to three calls at relatively low amplitude, with amplitude increasing with each call, after which amplitude remained relatively constant (Fig. 7). Individual calls decreased in amplitude over time (Fig. 7). Natural history: Megophrys fansipanensis sp. nov. is associated with upper montane bamboo forest, mixed bamboo forest and disturbed secondary broadleaf forest with a relatively open canopy. All individuals have been encountered at night and found in riparian habitat along 1–2 m wide streams with clear water and rocky stream beds. The majority of individuals were encountered in the month of June, fewer animals were observed in September. Males were calling from stream-side vegetation in June 2012 and June 2016. A gravid female was found adjacent to a fast-flowing stream in June 2016. Tadpoles were not observed at this time. During field work in September 2015, only a single unvouchered male was encountered, no vocalisations were heard and tadpoles were not observed. AMS R186115 VNMN 2018.01 AMS R186114 Number of call groups measured 5 3 12 Number of calls measured 20 20 20 Call duration (ms) 41.2 (38–45) 43.8 (41–49) 40.8 (34–45) Intercall interval (ms) 213.7 (197–245) 197.7 (180–227) 201.8 (168–290) Call repetition rate (calls/s) 3.8 4.0 4.0 Calls/call group 22.2 (18–25) 34.7 (24–41) 11.4 (1–22) Pulses/call 11.1 (10–12) 10.8 (10–11) 12.9 (12–14) Dominant frequency (kHz) 3.6 (na) 3.9 (3.9–4.1) 4.0 (3.6–4.7) Temperature (°C) 15.3 18.3 15.3 Distribution and conservation status: Megophrys fansipanensis sp. nov. is only known from Mount Fansipan at elevations between 2200 and 2813 m asl (Fig. 1). We did not detect this species during field work on Mount Fansipan at elevations of 1200 m asl (June 2016), 1700 m asl (June 2012 & 2016) and 1900 m asl (June 2016; September 2017). The high elevation sites where this species occurs face the immediate threat of habitat degradation and pollution due to the impact of tourists; habitats were noticeably polluted with garbage and runoff from toilets near the stream (Fig. 6D). In addition, if Megophrys fansipanensis sp. nov. is restricted to a narrow, high-elevation band, then the species is likely to be particularly vulnerable to climate change. Megophrys fansipanensis sp. nov. may occur more widely, but it was not detected in habitat similar to that found on Mount Fansipan during recent surveys on Mount Ky Quan San, 25 km north of our collection sites. The current EOO is predicted to be 630 km 2, qualifying Megophrys fansipanensis nov. sp. for listing as Endangered in accordance with the IUCN Red List of Threatened Species categories and criteria B1ab(iii). Comparisons: Megophrys fansipanensis sp. nov. can be distinguished from all species in the subgenus Panophrys found in mainland Southeast Asia, north of the Isthmus of Kra and neighbouring provinces of China on the basis of morphology, and from all species in the subgenus for which comparable data is available on the basis of molecular and acoustic data. Comparisons with each subgenus are discussed separately below. The following comparison is based on 13 males and two females of Megophrys fansipanensis sp. nov. Subgenus Brachytarsophrys: Megophrys fansipanensis sp. nov. can be distinguished from M. carinense, M. feae and M. intermedia by the absence of a transverse fold at the base of the head (versus presence in M. carinense, M. feae and M. intermedia) and having a smaller adult male size, SVL 30.9–44.3 mm (versus SVL> 79.1 mm; material examined). Subgenus Ophryophryne: Megophrys fansipanensis sp. nov. can be distinguished from all species in the subgenus Ophryophryne due to the presence (versus absence) of maxillary teeth (Poyarkov et al. 2017; material examined) and further from M. elfina by having a larger adult female size, SVL 41.7–42.5 mm (versus 35.1–36.5 mm, N =6, in M. elfina; Poyarkov et al. 2017) and the presence of vomerine teeth (versus absence in M. elfina; Poyarkov et al. 2017); from M. gerti by having a smaller adult female size, SVL 41.7–42.5 mm (versus 43.1–47.4 mm, N =3, in M. gerti; Poyarkov et al. 2017; material examined) and the presence of vomerine ridges and teeth (versus absence in M. gerti; Poyarkov et al. 2017); from M. hansi by having a smaller adult female size, SVL 41.7– 42.5 mm (versus 45.1–53.9 mm, N =5, in M. hansi; Poyarkov et al. 2017; material examined); from M. koui by the absence of a protruding fleshy projection above the cloaca in sexually mature males (secondary sexual characteristic of sexually mature M. koui; Poyarkov et al. 2017); and from M. synoria by having a smaller adult female size, SVL 41.7–42.5 mm (versus 51.4–70.7 mm, N =3, in M. synoria; Poyarkov et al. 2017) and the presence of microspinules on nuptial pads of sexually mature males (versus microgranules on nuptial pads of sexually mature male M. synoria; material examined). Subgenus Xenophrys: Megophrys fansipanensis sp. nov. differs from M. auralensis by having a smaller adult male size, SVL 30.9–44.3 mm (versus 60.1¯ 76.9 mm, N =20, in M. auralensis; Ohler et al. 2002; Neang et al. 2013); from M. damrei by having a smaller adult male size, SVL 30.9–44.3 mm (versus 47.7–57.1 mm, N =7, in M. damrei; Mahony 2011; Neang et al. 2013; material examined); from M. glandulosa by having a smaller adult male size, SVL 30.9–44.3 mm (versus 76.7–81.6 mm, N =10, in M. glandulosa; Fei et al. 2009; material examined), a lack of dermal fringes on toes (versus presence in M. glandulosa; Huang et al. 1998; Mahony et al. in press), a lack of distinct interdigital webbing between toes (versus basal webbing in M. glandulosa; material examined), and the absence of a light-coloured upper lip stripe (versus presence in M. glandulosa; Fei et al. 1990; Mahony et al. in press); from M. lekaguli by having a smaller adult male size, SVL 30.9–44.3 mm (versus 55.6–68.1 mm, N =8, in M. lekaguli; Stuart et al. 2006b; material examined); from M. major by having a smaller adult male size, SVL 30.9&nd

Megophrys (Panophrys) hoanglienensis sp. nov. Figs. 8���13 & Table 4. Holotype: One adult male (VNMN 2018.02), found beside a 5 m wide low-gradient shallow stream in slightly disturbed evergreen forest, Mount Fansipan, Hoang Lien National Park (HLNP), Sa Pa District, Lao Cai Province, Vietnam (22.33518�� N, 103.77947�� E, 2075 m asl; Fig. 8), collected at 19:15 h on 14 June 2012 by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. Paratypes: Four adult males (VNMN 0 7034, AMS R186118, AMS R186119 & AMS R186120), found in the vicinity of a 2 m wide low-gradient stream in disturbed evergreen forest, Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam (22.32827�� N, 103.78184�� E, 2242 m asl; Fig. 8), collected at 19:45 h on 12 June 2012; one adult female (AMS R186121), found beside a 5 m wide low-gradient shallow stream in slightly disturbed evergreen forest, Mount Fansipan, HLNP, Sa Pa District, Lao Cai Province, Vietnam (22.33518�� N, 103.77947�� E, 2075 m asl), collected at 19:15 h on 14 June 2012. All aforementioned specimens collected by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. Two adult males (AMS R186122 & AMS R186123), found calling from a rocky overhang 1.8 m above and 1.5 m away from a 3 m wide cascading stream at the base of Love Waterfall (th��c T��nh Y��u) in disturbed evergreen forest on Mount Fansipan, HLNP, Tam Duong District, Lai Chau Province, Vietnam (22.35065�� N, 103.77136�� E, 1898 m asl; Figs. 8 & 12 A���B), collected at 22:15 h on 24 June 2016 by Jodi J. L. Rowley, Benjamin Tapley, Cong Huy Le and Nguyen Thanh Chung; three adult males (AMS R186125, AMS R186126 & AMS R186127), found beside a swampy stream in disturbed evergreen forest, Mount Ky Quan San, Bat Xat District, Lao Cai Province, Vietnam (22.30343�� N, 103.37019�� E, 2153 m asl), collected at 21:00 h on 0 6 September 2017 by Nguyen Thanh Chung, Luan Thanh Nguyen, Luke Harding, Timothy Cutajar, Jodi J. L. Rowley and Benjamin Tapley. Referred specimens: One adult male (HLNP 20120624 00003), found calling from a rocky overhang 1.8 m above and 1.5 m away from a 3 m wide cascading stream at the base of Love Waterfall in disturbed evergreen forest on Mount Fansipan, HLNP, Tam Duong District, Lai Chau Province, Vietnam (22.35065�� N, 103.77136�� E, 1898 m asl; Figs. 8 & 12 A���B), collected at 22:15 h on 24 June 2016 by Jodi J. L. Rowley, Benjamin Tapley, Cong Huy Le and Nguyen Thanh Chung; two juvenile specimens (HLNP 20170906 0 0 0 0 4 & AMS R186124), found beside a 3 m wide stream in disturbed evergreen forest on Mount Fansipan, HLNP, Tam Duong District, Lai Chau Province, Vietnam (22.21039�� N, 103.46293�� E, 1882 m asl), collected at 20:00 h on 0 6 September 2017 by Nguyen Thanh Chung, Luan Thanh Nguyen, Luke Harding, Timothy Cutajar, Jodi J. L. Rowley and Benjamin Tapley; six adult males (HNUE MCC.2017.83���85, HNUE MCC.2017.88 & IEBR MCC.2017.86���87), found in the vicinity of Hang De Cha Hai stream, Che Tao Village, Che Tao Commune, Mu Cang Chai District, Mu Cang Chai Species Habitat Conservation Area, Yen Bai Province, Vietnam (21.76247�� N, 104.01889�� E, 2046 m asl), collected on 0 1 May 2017 by Yen Thi Do, Anh Ngoc Dao and Nam Hai Nguyen. Etymology: Specific epithet ��� hoanglienensis ��� is a toponym in reference to the type locality of the species, meaning ���from Hoang Lien��� Range. Suggested vernacular name: Hoang Lien horned frog (English), ���ch s���ng ho��ng li��n (Vietnamese). Diagnosis: Megophrys hoanglienensis sp. nov. differs from its congeners by a combination of the following morphological characters: (1) medium adult size, adult male SVL 37.4���47.6 mm (N= 11); adult female SVL 59.6 mm (N= 1); (2) small palpebral horn on upper eyelid present; (3) interdigital webbing and lateral fringes on the toes indistinct or absent; (4) snout mucronate in dorsal view; (5) TYD/EL 54.2���75.0% in adult males (N =11), 60.0% in adult female (N= 1); (6) subarticular tubercles absent; (7) palmar tubercles absent; (8) inner metatarsal tubercle present; (9) HW/SVL 37.4���42.5% in adult males (N= 11), 38.3% in adult female (N= 1); (10) SHL/SVL 44.3��� 62.5% in adult males (N =11), 57.5% in adult female (N= 1); (11) tympanum clearly defined; (12) TYE/SVL 6.3��� 9.5% in adult males (N= 11), 7.6% in adult female (N= 1); (13) vomerine ridges, vomerine and maxillary teeth present; (14) an advertisement call with a dominant frequency of 2.8���3.0 kHz at 18.5 ��C. Description of holotype (measurements provided in Table 4): Sexually mature male (Figs. 9 A���B, 10A���C, 11A���B). Head small, wider than long; snout mucronate in dorsal view, obtusely protruding in lateral view, rostral appendage absent; loreal region vertical and concave; canthus rostralis angular; dorsal region of the snout slightly concave; eye length 38.0% longer than the maximum tympanum diameter and subequal to the snout length; eye to tympanum distance greater than the maximum tympanum diameter; tympanum circular. Pupil in life oval, vertically orientated when dilated; nostrils orientated laterally, closer to the eyes than the snout; internarial distance subequal to the width of the eyelids, and subequal to the narrowest point between the upper eyelids; vomerine ridges present, oblique and barely separated anteriorly, with small vomerine teeth; maxillary teeth present. Tongue moderately large and feebly notched posteriorly. Forelimbs moderately long and thin; forearm length shorter than the hand length; fingers long and narrow without lateral fringes, finger length formula FI=FII=FIVF1 GURE 10. Megophrys hoanglienensis sp. nov. in preservative (holotype VNMN 2018.02). (A) Lateral view of head, (B) palmar surface of right hand, and (C) plantar surface of left foot. 5 mm scale bar. Skin of dorsal surfaces of the body, limbs, and both dorsal and lateral surfaces of the head weakly granular; tympanum granular with borders slightly raised; very small palpebral horn present on the outer edge of the upper eyelid; supratympanic fold narrows as it passes above the tympanum, widening again before terminating above the axilla; flanks with small scattered tubercles; thin dorsolateral fold extending posteriorly from behind the supratympanic fold to the groin on each side; a weak, ���V���-shaped parietoscapular ridge present, its two sides extending posteriorly from above the tympanum and meeting medially beyond the level of the axilla; a second weak inverted ���U���-shaped sacral ridge present on the mid-dorsum which joins laterally with the dorsolateral folds; ���V��� and ���U���-shaped ridges not joined at their apex, apex of ridges with black asperities; small tubercles present on the dorsal surface of the thighs, shanks and forearms. Large distinct tubercles present on the flanks, inguinal region and dorsal surface of the hindlimbs; chest and ventral surfaces of the forelimbs and shanks smooth; pectoral glands obvious, small, slightly raised, positioned level with the axilla; femoral glands small, slightly raised, one positioned equidistant from the knee and the cloaca on the posterior surface of each thigh; white skin asperities forming narrow band circummarginally on the upper and lower jaw; black asperities on the posterior parts of the upper eyelids, on the dorsum and area directly above the vent, and on tubercles of the tympanic regions, absent elsewhere. Colour of holotype: In life (Figs. 11 A���B): Dorsally dark tan; with light tan blotches; darker brown triangular marking between the eyes with a lighter central blotch; cream supratympanic folds with black lower margin; inguinal regions light tan; dark brown hindlimbs with darker blotches; posterior surface of the thighs and shanks very dark brown; forearms light brown with a single dark bar; dorsal surface of the fingers with dark brown bars; gular, pectoral and abdomen region mottled grey-brown and cream, gular region with a medial dark brown stripe; ventral surface of thighs unmarked; pectoral glands off-yellow; iris metallic gold. In preservative (Figs. 9 A���B & 10A���C): Markings are the same as in life, colouration differences are as follows: The majority of the dorsal and lateral surfaces of the head, body, forelimbs and hindlimbs brown; dorsolateral and supratympanic folds and flank tubercles brownish-cream; tip of the snout cream, lateral canthus rostralis dark brown; wide vertical dark brown bar below the eyes; dark brown band running from the posterior edge of the eyes, through the tympanum, and terminating where the forelimbs join the body; lower margin of the supratympanic folds dark brown, sharply contrasting with creamy grey upper margin; gular region, chest and anterior part of the abdomen primarily creamy-grey with brown speckling, and a dark brown medial bar in the gular region; dark brown blotches anterior to each of the pectoral glands, pectoral glands light cream in colour; abdomen speckled with dark brown; ventral surfaces of the thighs and shanks with diffuse pale brown mottling; ventral surfaces of the feet grey brown with cream digit tips; area surrounding the vent dark brown; forelimbs ventrally mottled with dark brown bar running longitudinally on the outer ventral surface of the forearms; grey colouration of ventral side of the hands with darker mottling and cream digit tips; femoral glands cream. Variation: Morphometric measurements of the type series and referred specimens are provided in Table 4. Nuptial pads were not visible on one male specimen (AMS VNMN 07034) although sex was confirmed via inspection of the gonads. Two (AMS R186120 & AMS R186121) of the 12 specimens possessed rudimentary webbing between the toes. Four (VNMN 0 7034 & AMS R186118��20) of the 12 specimens with indistinct lateral fringes on the toes. It is not clear if the presence of indistinct fringes on the toes of these specimens is an artefact of the positioning during the fixation process. Relative finger lengths are highly variable, and only one specimen (AMS R186119; FI���FIISecondary sexual characters: All sexually mature males had slightly raised nuptial pads covered with black microspinules, nuptial pads covering most of the dorsal surface of the base of FI and FII; nuptial pad oval shaped on FI and FII; the protruding fleshy projection posterior to the cloaca (a secondary sexual character of some male Megophrys, e.g., M. caudoprocta, M. koui and M. pachyproctus) is absent on all male specimens. The examined female did not possess nuptial pads. Molecular data: Sequences generated from nine adult specimens in this study (Table 1) were most similar to ��� Megophrys sp. 5��� (sensu Chen et al. 2017) collected in Sa Pa District, Lao Cai Province, Vietnam; 16S rDNA sequences differed by only 0.3% uncorrected p -distance between ��� Megophrys sp. 5��� and our specimens (from Lao Cai & Lai Chau provinces), and by 1.1% for specimens collected approximately 58 km away in Yen Bai Province, which is greater than the intraspecific variation reported elsewhere within the subgenus Panophrys (e.g., Li et al. 2014 reported an intraspecific p -distance of 0.6% between populations of M. obesa Wang, Li & Zhao, 2014), but is lower than intraspecific variation within Megophrys sensu lato (e.g., 3.1% mean uncorrected p -distance in M. elfina; Poyarkov et al. 2017). Uncorrected p -distance was 0.8% between specimens of Megophrys hoanglienensis sp. nov. collected from Lao Cai and Lai Chau provinces. The smallest interspecific p- distance in our study was between Megophrys hoanglienensis sp. nov. and Megophrys fansipanensis sp. nov. at 3.3% where the two new species occur in syntopy supporting their position as distinct sister taxa. Advertisement call: Call descriptions are based on the calls of one paratype (AMS R186122), five call groups and 20 calls in total were analysed. Advertisement calls were recorded at 18.5 ��C ambient temperature. Calls were an average of 102.9 ms (96���108 ms) in duration (Table 5; Fig. 13). The average number of pulses per call was 18.7 (12���22). The average dominant frequency of all the analysed calls was 3.0 kHz (2.8���3.0 kHz). Calls were repeated at a rate of 2.6 calls per second for the call groups analysed, and had an average intercall interval of 274.2 ms (178��� 565 ms). The number of calls within each call group ranged from 11 to 21, with an average of 17. Most call groups began at a relatively low amplitude, increasing with each call up to approximately a quarter of the duration, after which amplitude varied but did not follow a trend or pattern (Fig. 13A & B). Individual calls began with a medium relative amplitude but peaked early before declining towards the end, and the interval between the first and second pulse was longer than those between any of the following pulses (Fig. 13C). Natural history: Megophrys hoanglienensis sp. nov. is associated with disturbed secondary broadleaf forest with a relatively open canopy. All individuals have been found at night in riparian habitat along clear water, 2���5 m wide streams with a gravel or rocky substrate. Males were calling from streamside vegetation in June 2012 and June 2016 but not during the surveys undertaken in May or September 2017. A non-gravid female was found adjacent to a wide, low-gradient, shallow stony stream in June 2016. Tadpoles were not observed at any of the locations where this species has been found (surveys undertaken in June 2012 and 2016, and in September 2015 and 2017). Megophrys hoanglienensis sp. nov. is sympatric with M. jingdongensis at 1990 m asl on Mount Fansipan, and with M. fansipanensis sp. nov. on Mount Fansipan at 2242 m asl. Megophrys hoanglienensis sp. nov. is sympatric with M. jingdongensis and M. rubrimera on Mount Ky Quan San at 2153 m asl (Tapley et al. 2018b). Distribution and conservation status: This species is known from between 1898���2242 m asl at four localities, up to 92 km apart in the Hoang Lien Range (Sa Pa District, Lao Cai Province, HLNP; Tam Duong District, Lai Chau Province; Mount Ky Quan San, Bat Xat District, Lao Cai Province and Che Tao Commune, Yen Bai Province, Vietnam). The species��� EOO is currently predicted to be 3216 km 2 (Fig. 8). Ongoing disturbance to the species��� habitat due to forest clearance for agriculture has been observed in the parts of Sa Pa District where the species was observed. In Bat Xat District the forest in which this species occurs is being negatively impacted by fuelwood collection for the tourism industry and by the grazing of livestock. Megophrys hoanglienensis sp. nov. is likely to qualify as Endangered in accordance with the IUCN Red List of Threatened Species categories and criteria B1ab(iii). Comparisons: Megophrys hoanglienensis sp. nov. can be distinguished from all species in the subgenus Panophrys found in mainland Southeast Asia, north of the Isthmus of Kra and neighbouring provinces of China on the basis of morphology, and from all species in the subgenus for which comparable data is available on the basis of molecular and acoustic data. Comparisons with each subgenus are discussed separately below. The following comparison is based on 11 males and one female of Megophrys hoanglienensis sp. nov. Subgenus Brachytarsophrys: Megophrys hoanglienensis sp. nov. can be distinguished from M. carinense, M. feae and M. intermedia by the absence of a transverse fold at the base of the head (versus presence in M. carinense, M. feae and M. intermedia), and by having a smaller adult male size, SVL 41.1���47.6 mm (versus SVL> 79.1 mm; material examined). Subgenus Ophryophryne: Megophrys hoanglienensis sp. nov. can be distinguished from all species in the subgenus Ophryophryne due to the presence (versus absence) of maxillary teeth (Poyarkov et al. 2017; material examined); further from M. elfina by having a larger adult female size, SVL 59.6 mm (versus 35.1���36.5 mm, N =6, in M. elfina; Poyarkov et al. 2017); from M. gerti by having a larger adult female size, SVL 59.6 mm (versus 43.1��� 47.4 mm, N =3, in M. gerti; Poyarkov et al. 2017); from M. hansi by having a larger adult female size, SVL 59.6 mm (versus 45.1���53.9 mm, N =5, in M. hansi; Poyarkov et al. 2017; material examined); from M. koui by the absence of a protruding fleshy projection above the cloaca in sexually mature males (secondary sexual characteristic of adult M. koui; Poyarkov et al. 2017); and from M. synoria by the presence of microspinules on nuptial pads in adult males (versus microgranules on nuptial pads in adult M. synoria; Poyarkov et al. 2017; material examined). Subgenus Xenophrys: Megophrys hoanglienensis sp. nov. differs from M. auralensis by having a smaller adult male body size, SVL 37.4���47.6 mm (versus 60.1���76.9 mm, N = 20 in M. auralensis, N =20; Ohler et al. 2002; Neang et al. 2013); from M. damrei by having a smaller adult female body size, SVL 59.6 mm (versus 69.1 mm, N= 1, in M. damrei; Mahony 2011; material examined); from M. glandulosa by having a smaller adult male body size, SVL 37.4���47.6 mm (versus 76.7���81.6 mm, N =10, in M. glandulosa; Fei et al. 2009; Fei et al. 1990; material examined), absence of distinct lateral fringes on toes (versus presence in M., Published as part of Tapley, Benjamin, Cutajar, Timothy, Mahony, Stephen, Nguyen, Chung Thanh, Dau, Vinh Quang, Luong, Anh Mai, Le, Dzung Trung, Nguyen, Tao Thien, Nguyen, Truong Quang, Portway, Christopher, Luong, Hao Van & Rowley, Jodi J. 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Kane, Daniel, Tapley, Benjamin (2018): Description and development of the tadpole of Rhacophorus feae (Anura; Rhacophoridae). Zootaxa 4504 (1): 138-144, DOI: https://doi.org/10.11646/zootaxa.4504.1.9

Rowley, Jodi J. L., Tapley, Benjamin, Chung, Nguyen Thanh, Altig, Ronald (2017): Tadpole of the Critically Endangered Sterling's Toothed Toad (Oreolalax sterlingae). Zootaxa 4272 (4): 579-582, DOI: https://doi.org/10.11646/zootaxa.4272.4.6

Megophrys (Panophrys) rubrimera sp. nov. Figs. 2, 3, 4, 5 & 6; Table 3. –– Megophrys kuatunensis Orlov et al. 2000:10 (partim: Megophrys cf. kuatunensis “Mount Fan Si Pan, Hoang Lien Range (Sa Pa District, Lao Cai Province)”; Orlov et al. 2002:83 (partim: “northern Vietnam”); Nguyen et al. 2009:88 (partim: Sa Pa District, Lao Cai Province, Vietnam). Holotype: VNMN 2017.002 adult male calling beside a 1 m wide rocky stream (stream bed 5–6 m wide) in heavily disturbed evergreen forest, Sa Pa District, Lao Cai Province, Vietnam (22.38205°N, 103.78745°E, 1708 m asl; Fig. 1). Collected at 22:15 h on 18 June 2012 by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. Paratypes: Two male specimens (VNMN 2017.003, AMS R177676) collected from a 3.5 m wide rocky stream in slightly disturbed evergreen forest, Sa Pa District, Lao Cai Province, Vietnam (22.39829°N, 103.78545°E, 1400 m asl; Fig. 1) on 17 June 2012 by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. One adult male specimen ( AMS R177675) collected from disturbed habitat; a 2 m wide rocky stream in slightly disturbed evergreen forest, Sa Pa District, Lao Cai Province, Vietnam (22.38208°N, 103.78699°E, 1722 m asl) between 22:00 and 22:30 h on 22 June 2016 by Jodi J. L. Rowley, Benjamin Tapley and Nguyen Thanh Chung. Three adult male specimens ( AMS R177677, AMS R177678 and AMS R177679) collected from disturbed habitat; a steep north-easterly facing road side bank with seepages and a small stream, with evergreen forest nearby, Sa Pa District, Lao Cai Province, Vietnam (22.3809°N, 103.78798°E, 1714 m asl; Figs 1 & 7 C–E) between 22:00 and 22:30 h on 23 June 2016 by Jodi J. L. Rowley, Benjamin Tapley and Nguyen Thanh Chung. Referred specimens: One adult male specimen (field tag JJLR03813 - HNLP2016062200001) collected from disturbed habitat; a 2 m wide rocky stream in slightly disturbed evergreen forest, Sa Pa District, Lao Cai Province, Vietnam (22.38208°N, 103.78699°E, 1722 m asl) between 22:00 and 22:30 h on 22 June 2016 by Jodi J. L. Rowley, Benjamin Tapley and Nguyen Thanh Chung. Specimen deposited at Hoang Lien National Park Headquarters, Vietnam. Tadpole specimen (AMS R177680) collected from a pool in a 2 m wide rocky stream in slightly disturbed evergreen forest, Sa Pa District, Lao Cai Province, Vietnam (22.38208°N, 103.78699°E, 1722 m asl) between 22:00 and 22:30 h on 22 June 2016 by Jodi J. L. Rowley, Benjamin Tapley and Nguyen Thanh Chung. The pH of the water body was 7.7, alkalinity 0.1 ppm and the water temperature was 17.6 °C. Etymology: The specific name “ rubrimera ”, an invariable noun in apposition, derived from the Latin word ruber (prefix rubri-) meaning red, and the Latinised version of the Greek noun mera, meaning thigh, in reference to the bright red-orange colouration of the groin, and the inner and outer thighs of the new species. Suggested vernacular name: Red-thighed horned frog (English), Cóc sừng đùi đỏ (Vietnamese) Diagnosis: Assigned to the genus Megophrys on the basis of tadpole morphology; tadpoles have dorsally orientated umbelliform oral discs (Dubois & Ohler 1998; Li et al. 2011), which is diagnostic for the genus within Megophryidae; and to the subgenus Panophrys on the basis of molecular data. Megophrys rubrimera sp. nov. differs from its congeners by a combination of the following morphological characters (based on eight adult males): (1) small size (SVL 26.7–30.5 mm); (2) very small palpebral horn on upper eyelid; (3) toes lacking interdigital webbing but possessing narrow lateral fringes; (4) tympanum diameter: eye diameter 58.0–76.0 mm; (5) shank length: snout vent length 48.0–56.0%; (6) groin, inner surface of thighs and outer surface of shanks red-orange; (7) absence of subarticular tubercles on fingers and toes; (8) red-orange inner metatarsal tubercle; (9) head width greater than head length; (10) head width: snout vent length 38.0–42.0%; (11) weakly defined vomerine ridges with teeth; and (12) an advertisement call with a dominant frequency of 3.23 kHz. Description of holotype: Sexually mature male (Figs. 2 & 3). Head small, wider than long; the snout rounded in dorsal view and obtusely protruding in lateral view, rostral appendage absent (Figs. 3A & 4G); loreal region vertical and concave; canthus rostralis angular; dorsal region of the snout is slightly concave; eye diameter nearly twice the maximum tympanum diameter and subequal to snout length; eye-tympanum distance is smaller than the maximum tympanum diameter; tympanum round and orientated vertically. Pupil in life oval, vertically orientated when dilated; nostril orientated laterally, closer to eye than snout; internarial distance exceeds eyelid width, and subequal to narrowest point between upper eyelids; weakly defined vomerine ridges present, oblique and barely separated anteriorly with small vomerine teeth; maxillary teeth present. Tongue moderately large and not clearly notched posteriorly. Forelimbs short and stocky, forearm length shorter than hand length; fingers short and wide without lateral fringes (Fig. 3B), finger length formula I Skin of dorsal surfaces of body, limbs, and dorsal and lateral surfaces of head weakly granular; tympanum granular with its borders slightly raised; very small pointed tubercle present on outer edge of upper eyelid; supratympanic fold narrows as it passes above the tympanum, terminating above axilla, supratympanic fold with many dark asperities along its crest (Fig. 4A); flanks with small scattered tubercles, some of which terminate in darkened asperities; thin dorsolateral fold extending from behind supratympanic fold to approximately threequarters distance to groin; a weak, “V”-shaped parietoscapular ridge present, its two sides extending posteriorly from above tympanum and meeting medially beyond level of axilla; a second inverted “U”-shaped ridge present on mid-dorsum which joins laterally with dorsolateral folds (Figs. 2A & 4A); ridges with some dark asperities; small tubercles tipped with dark asperities on dorsal surface of shanks, and arranged into distinct transverse rows on the thighs and forearms. Large distinct tubercles present in the inguinal region; gular region, chest and ventral surfaces of limbs smooth; pectoral glands obvious, small, slightly raised, positioned on level with axilla (Figs. 2B & Fig. 4D); femoral glands small, slightly raised, one positioned equidistant from knee and cloaca on the posterior surface of each thigh. Colour of holotype in life (Figs. 4A–B, D & F): Dorsally light tan; darker “Y”-shaped marking on dorsum between the eyes, border of marking beige; vertical dark brown bar below eyes; lateral surface of the eyelid dark with a single medial light bar; tympanum dark brown; hind and forelimbs possess dark tan bars, darker on the forelimbs than the hind limbs; tubercles on flanks encircled by darker tan; groin red-orange; gular and pectoral region grey-brown; abdomen light grey with dark grey blotches and white speckling; ventral surface of thighs pinkish grey with darker blotches; inner thighs and outer surface of shanks red-orange; underside of Fingers III, IV orange; thenar tubercles red-orange; inner metatarsal tubercle on feet red-orange; iris metallic brown. Colour of holotype in preservative: ( Figs. 2 & 3): Majority of dorsal and lateral surfaces of the head, body, forelimbs and hind limbs brown; darker brown “Y”-shaped marking between eyes; darker brown “X”-shaped marking over opposing “V”-shaped and “U”-shaped ridges; dorsolateral folds and flank tubercles bordered by darker margin; front of snout and lateral canthus rostralis dark brown; narrow vertical dark brown bar below eyes and dark brown blotch covering tympanum; upper jaw with darker blotches; two dark brown blotches on the dorsal surface of forearms; dorsal surface of Fingers II, III and IV with dark brown blotches. Gular region, chest and anterior part of abdomen primarily creamy-grey, with grey brown speckling on gular region; abdomen blotched with dark brown; ventral surfaces of thighs with grey brown mottling, and shanks with grey brown speckling; ventral surfaces of feet grey brown; area surrounding vent with dark brown blotches; forelimbs ventrally mottled and blotched with light and dark brown; extending to the ventral surface of hands; inner metatarsal tubercles unmarked and beige, tips of fingers unmarked and beige; tips of toes only lightly speckled with grey brown, if at all; lateral fringe on toes, beige, unmarked; pectoral and femoral glands beige. Measurements (in mm): Holotype. SVL 29.8, HW 11.2, HL 9.4, SL 3.9, SN 2.0, EN 1.6, EL 3.3, IUE 4.0, UEW 3.1, IFE 3.5, IBE 3.8, TYD 2.0, TYE 1.9, FAL 6.5, HAL 7.5, FIL 2.4, FIIL 2.8, FIIIL 4.1, FIVL 2.9, TL 14.6, SHL 14.4, FOL 13.5, IMT 2.7, weight in life 2.4 g. Variation: Measurements of the type series are shown in Table 3. Paratypes and referred specimens generally agree with the holotype morphologically, but with the following exceptions: in VNMN 2017.003, AMS R177676 the middorsal “U”-shaped ridge (on the holotype) is replaced by an inverted “V”-shaped ridge. AMS R177676 with distinct dark grey blotches on ventral surface of thighs, this is lacking in VNMN 2017.003 and the holotype specimen. In life, AMS R177676 has a vivid red-orange flash on the axillary region (Fig. 4G). Gular region of VNMN 2017.002 and AMS R177678 lack dark blotches, present in all other specimens. Ventral surface of thighs (on AMS R177679 and AMS R177676) are covered in dark blotches, which are not present on any of the other specimens. Tongue weakly notched in one specimen (HNLP 2016062200001). Secondary sexual characters: Females of Megophrys rubrimera sp. nov. are currently unknown. Two male specimens (AMS R177678 and AMS R177679) possessed raised oval nuptial pads covered in microspinules on the base of Finger II; protruding fleshy projection (secondary sexual characteristic of some male Megophrys e.g. M. caudoprocta, M. koui and M. pachyproctus) above the cloaca absent from all specimens. After fixation it was very difficult to open the mouths of the specimens without damaging them due to the small head size. It was therefore not possible to determine the position, presence or absence of internal vocal slits which are a secondary sexual characteristic used as a diagnostic character to the species level (e.g. Mahony et al. 2013). Tadpole: (Fig. 5): The following tadpole description is based on a single specimen (AMS R177680) at Stage 37, the tadpole was confirmed as that of M. rubrimera sp. nov. by molecular analysis. Body elongated and slender; nares oval and are closer to the eye than to the snout; internarial distance is subequal to the interorbital distance; eyes positioned dorsolaterally, the pupils are round; the spiracle is sinistral and the spiracular tube protrudes from midway along the body wall just posterior to maximal trunk width and opens laterally; the tail makes up 68% of the total body length; the dorsal tail fin is low, particularly at the proximal half of the tail length; the basal tail width is 60% of the maximal trunk width; the oral disk is subterminal and antero-dorsal; the width of the umbelliform oral disc makes up ~80% of the maximal width of the trunk; the upper and lower lips each have deep medial emarginations; submarginal papillae are present on both sides between the oral orifice and the emarginations of the upper and lower lip, concentrated in a row around the margins; keratodonts are absent. Colour in life: Dorsally, the head and body are brown with darker speckles; obvious pale neuromasts; the oral disk is translucent yellowish brown with dark brown submarginal papillae; the dorsal and ventral fins opaque, pale yellowish brown with speckles; the venter is speckled white and brown; iris orange, speckled with black dots. Colour in preservative: Body brown with darker speckles; oral disc translucent grey brown with dark brown submarginal papillae; the dorsal and ventral fins opaque, pale grey brown with darker speckles; the venter is speckled grey and brown. Tadpole body measurements (in mm): BL 10.5, BH 4.4, BS 4.3, ES 1.7, IND 1.7, IOD 4.1, BW 5.4, ED 1.2, LFH 1.3, MTH 5.9, NE 1.9, ODW 4.5, SN 0.6, SS 5.7, TAL 22.8, TTL 33.3, TMH 3.1, TMW 5.4, UFH 1.3). Advertisement call: Call descriptions are based on the calls of one paratype (AMS R177677) and two unvouchered individuals. Advertisement calls were recorded at 21.0–22.9 °C ambient temperature. Calls were an average of 73.3 ms (62–85 ms) in duration (Table 4; Fig. 6). Calls of unvouchered individual (b) contained an average of 23.05 pulses (19–25), whereas the calls of AMS R177677 and another unvouchered individual (a) were not distinctly pulsed. The average dominant frequency of calls was 3.3 kHz (3.2–3.4 kHz). Calls were repeated at a rate of approximately 3.25 (3.05–3.37) calls per second, and had an average intercall interval of 221.3 ms (190–261 ms). The number of calls within each call group ranged from 16–51, with an average of 38.7. Most call groups began at a relatively low amplitude, increasing with each call up to approximately a quarter to a third of the duration, after which amplitude remained relatively constant (Fig. 6A i –ii & B i–ii). Individual calls either began with a medium relative amplitude and peaked near the middle before declining towards the end (Fig. 6A iii), or peaked at the beginning, steadily declining throughout (Fig. 6B iii). Natural history: All specimens of Megophrys rubrimera sp. nov. were found in disturbed evergreen secondary forest. Males were calling during June 2012 and 2016, and tadpoles (Stage 37) were collected in June 2016, suggesting a prolonged breeding season or that the species has a long larval period. Males were observed calling on stream-side vegetation (Fig. 7A–B) and a steep north-eastern facing roadside bank with seepages and a small stream (Fig. 7C–E). Females were not observed. Distribution and conservation status: This species is known from between 1400 m asl and 1722 m asl at two localities, 2 km apart in Sa Pa District, Lao Cai Province, Vietnam and 50 km north-west at Maandi, Jinping County, Ailao Mountain Range, Yunnan Province, China (Fig. 1; Chen et al. 2017). The species' area of occupancy (AOO) and extent of occurrence (EOO) are currently predicted to be 385 km 2 and 2298 km 2, respectively. The Vietnamese and Chinese portions of this species’ range comprise two threat-defined locations; land use likely differs between the two countries, and habitat loss probably affects the species independently in each. Ongoing disturbance to the species’ habitat due to forest clearance for agriculture has been observed in Sa Pa District. We recommend that Megophrys rubrimera sp. nov. is listed as Endangered in accordance with the IUCN Red List of Threatened Species categories and criteria B1ab(iii). Comparisons: Megophrys (Panophrys) rubrimera sp. nov. can be distinguished from all other congeners found in mainland southeast Asia, north of the Isthmus of Kra and neighbouring provinces of China on the basis of morphology, and from all congeners for which comparable data is available on the basis of molecular and acoustic data. Comparisons with each subgenus are discussed separately below. Subgenus Brachytarsophrys: Megophrys rubrimera sp. nov. can be distinguished from the species in the subgenus Brachytarsophrys (M. carinense, M. feae and M. intermedia) in mainland southeast Asia and neighbouring provinces of China by the absence of a transverse fold at the base of the head (versus presence), and having a smaller adult male size, SVL 26.7–30.5 mm (versus SVL> 79.1 mm; examined material). Subgenus Ophryophryne: Megophrys rubrimera sp. nov. can be distinguished from M. elfina by the presence of lateral fringes on the toes (versus absence; Poyarkov et al. 2017) and by the presence of vomerine teeth (versus absence Poyarkov et al. 2017); from M. gerti by having a smaller adult male size, SVL 26.7–30.5 mm (versus 31.7–42.2 mm; Poyarkov et al. 2017; material examined); from M. hansi by having a smaller adult male size, SVL 33.4–43.1 mm; Ohler 2003; Poyarkov et al. 2017; material examined); from M. microstoma by having a smaller adult male size, SVL 26.7–30.5 mm (versus 34.3–44.4 mm; Poyarkov et al. 2017; material examined; from M. koui by the absence of a protruding fleshy projection above the cloaca in sexually mature males (secondary sexual characteristic of sexually mature M. koui); from M. synoria by having a smaller adult male size, SVL 26.7–30.5 mm (versus 38.2–53.7 mm; Stuart et al. 2006b; Poyarkov et al. 2017; material examined) and further from all species in the subgenus Ophryophryne due to the presence (versus absence) of maxillary teeth. FFIGURE 7. Megophrys rubrimera sp. nov. in situ, and habitat at collection localities. (A) Adult male paratype AMS R177675 and (B) unvouchered calling male. (C–E) Habitat at type locality in Sa Pa District, Lao Cai Province, Vietnam. Subgenus Xenophrys: Megophrys rubrimera sp. nov. differs from M. auralensis by having a smaller adult male size, SVL 26.7–30.5 mm (versus 60.1–76.7 mm; Ohler et al. 2002; Neang et al. 2013), and a lack of interdigital toe webbing (versus rudimentary webbing); from M. damrei by having a smaller adult male size, SVL 26.7–30.5 mm (versus 47.5–57.1 mm: Mahony 2011; Neang et al. 2013; material examined), a lack of interdigital toe webbing (versus rudimentary), and the presence of lateral fringes on the toes (versus absence); from M. glandulosa by having a smaller adult male size, SVL 26.7–30.5 mm (versus 76.7–81.6 mm: Fei et al. 2009; material examined), and a lack of interdigital webbing between the toes (versus basal webbing); from M. lekaguli by having a smaller adult male size, SVL 26.7–30.5 mm (versus 55.6–68.1

Megophrys (Panophrys) rubrimera sp. nov. Figs. 2, 3, 4, 5 & 6; Table 3. ������ Megophrys kuatunensis Orlov et al. 2000:10 (partim: Megophrys cf. kuatunensis ���Mount Fan Si Pan, Hoang Lien Range (Sa Pa District, Lao Cai Province)���; Orlov et al. 2002:83 (partim: ���northern Vietnam���); Nguyen et al. 2009:88 (partim: Sa Pa District, Lao Cai Province, Vietnam). Holotype: VNMN 2017.002 adult male calling beside a 1 m wide rocky stream (stream bed 5���6 m wide) in heavily disturbed evergreen forest, Sa Pa District, Lao Cai Province, Vietnam (22.38205��N, 103.78745��E, 1708 m asl; Fig. 1). Collected at 22:15 h on 18 June 2012 by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. Paratypes: Two male specimens (VNMN 2017.003, AMS R177676) collected from a 3.5 m wide rocky stream in slightly disturbed evergreen forest, Sa Pa District, Lao Cai Province, Vietnam (22.39829��N, 103.78545��E, 1400 m asl; Fig. 1) on 17 June 2012 by Jodi J. L. Rowley, Dau Quang Vinh, Pham Van Sang, Tran Van Tu, Hang A Su, Hoang A Di and Dinh Van Xuan. One adult male specimen ( AMS R177675) collected from disturbed habitat; a 2 m wide rocky stream in slightly disturbed evergreen forest, Sa Pa District, Lao Cai Province, Vietnam (22.38208��N, 103.78699��E, 1722 m asl) between 22:00 and 22:30 h on 22 June 2016 by Jodi J. L. Rowley, Benjamin Tapley and Nguyen Thanh Chung. Three adult male specimens ( AMS R177677, AMS R177678 and AMS R177679) collected from disturbed habitat; a steep north-easterly facing road side bank with seepages and a small stream, with evergreen forest nearby, Sa Pa District, Lao Cai Province, Vietnam (22.3809��N, 103.78798��E, 1714 m asl; Figs 1 & 7 C���E) between 22:00 and 22:30 h on 23 June 2016 by Jodi J. L. Rowley, Benjamin Tapley and Nguyen Thanh Chung. Referred specimens: One adult male specimen (field tag JJLR03813 - HNLP2016062200001) collected from disturbed habitat; a 2 m wide rocky stream in slightly disturbed evergreen forest, Sa Pa District, Lao Cai Province, Vietnam (22.38208��N, 103.78699��E, 1722 m asl) between 22:00 and 22:30 h on 22 June 2016 by Jodi J. L. Rowley, Benjamin Tapley and Nguyen Thanh Chung. Specimen deposited at Hoang Lien National Park Headquarters, Vietnam. Tadpole specimen (AMS R177680) collected from a pool in a 2 m wide rocky stream in slightly disturbed evergreen forest, Sa Pa District, Lao Cai Province, Vietnam (22.38208��N, 103.78699��E, 1722 m asl) between 22:00 and 22:30 h on 22 June 2016 by Jodi J. L. Rowley, Benjamin Tapley and Nguyen Thanh Chung. The pH of the water body was 7.7, alkalinity 0.1 ppm and the water temperature was 17.6 ��C. Etymology: The specific name ��� rubrimera ���, an invariable noun in apposition, derived from the Latin word ruber (prefix rubri-) meaning red, and the Latinised version of the Greek noun mera, meaning thigh, in reference to the bright red-orange colouration of the groin, and the inner and outer thighs of the new species. Suggested vernacular name: Red-thighed horned frog (English), C��c s���ng đ��i đ��� (Vietnamese) Diagnosis: Assigned to the genus Megophrys on the basis of tadpole morphology; tadpoles have dorsally orientated umbelliform oral discs (Dubois & Ohler 1998; Li et al. 2011), which is diagnostic for the genus within Megophryidae; and to the subgenus Panophrys on the basis of molecular data. Megophrys rubrimera sp. nov. differs from its congeners by a combination of the following morphological characters (based on eight adult males): (1) small size (SVL 26.7���30.5 mm); (2) very small palpebral horn on upper eyelid; (3) toes lacking interdigital webbing but possessing narrow lateral fringes; (4) tympanum diameter: eye diameter 58.0���76.0 mm; (5) shank length: snout vent length 48.0���56.0%; (6) groin, inner surface of thighs and outer surface of shanks red-orange; (7) absence of subarticular tubercles on fingers and toes; (8) red-orange inner metatarsal tubercle; (9) head width greater than head length; (10) head width: snout vent length 38.0���42.0%; (11) weakly defined vomerine ridges with teeth; and (12) an advertisement call with a dominant frequency of 3.23 kHz. Description of holotype: Sexually mature male (Figs. 2 & 3). Head small, wider than long; the snout rounded in dorsal view and obtusely protruding in lateral view, rostral appendage absent (Figs. 3A & 4G); loreal region vertical and concave; canthus rostralis angular; dorsal region of the snout is slightly concave; eye diameter nearly twice the maximum tympanum diameter and subequal to snout length; eye-tympanum distance is smaller than the maximum tympanum diameter; tympanum round and orientated vertically. Pupil in life oval, vertically orientated when dilated; nostril orientated laterally, closer to eye than snout; internarial distance exceeds eyelid width, and subequal to narrowest point between upper eyelids; weakly defined vomerine ridges present, oblique and barely separated anteriorly with small vomerine teeth; maxillary teeth present. Tongue moderately large and not clearly notched posteriorly. Forelimbs short and stocky, forearm length shorter than hand length; fingers short and wide without lateral fringes (Fig. 3B), finger length formula I Colour of holotype in life (Figs. 4A���B, D & F): Dorsally light tan; darker ���Y���-shaped marking on dorsum between the eyes, border of marking beige; vertical dark brown bar below eyes; lateral surface of the eyelid dark with a single medial light bar; tympanum dark brown; hind and forelimbs possess dark tan bars, darker on the forelimbs than the hind limbs; tubercles on flanks encircled by darker tan; groin red-orange; gular and pectoral region grey-brown; abdomen light grey with dark grey blotches and white speckling; ventral surface of thighs pinkish grey with darker blotches; inner thighs and outer surface of shanks red-orange; underside of Fingers III, IV orange; thenar tubercles red-orange; inner metatarsal tubercle on feet red-orange; iris metallic brown. Colour of holotype in preservative: ( Figs. 2 & 3): Majority of dorsal and lateral surfaces of the head, body, forelimbs and hind limbs brown; darker brown ���Y���-shaped marking between eyes; darker brown ���X���-shaped marking over opposing ���V���-shaped and ���U���-shaped ridges; dorsolateral folds and flank tubercles bordered by darker margin; front of snout and lateral canthus rostralis dark brown; narrow vertical dark brown bar below eyes and dark brown blotch covering tympanum; upper jaw with darker blotches; two dark brown blotches on the dorsal surface of forearms; dorsal surface of Fingers II, III and IV with dark brown blotches. Gular region, chest and anterior part of abdomen primarily creamy-grey, with grey brown speckling on gular region; abdomen blotched with dark brown; ventral surfaces of thighs with grey brown mottling, and shanks with grey brown speckling; ventral surfaces of feet grey brown; area surrounding vent with dark brown blotches; forelimbs ventrally mottled and blotched with light and dark brown; extending to the ventral surface of hands; inner metatarsal tubercles unmarked and beige, tips of fingers unmarked and beige; tips of toes only lightly speckled with grey brown, if at all; lateral fringe on toes, beige, unmarked; pectoral and femoral glands beige. Measurements (in mm): Holotype. SVL 29.8, HW 11.2, HL 9.4, SL 3.9, SN 2.0, EN 1.6, EL 3.3, IUE 4.0, UEW 3.1, IFE 3.5, IBE 3.8, TYD 2.0, TYE 1.9, FAL 6.5, HAL 7.5, FIL 2.4, FIIL 2.8, FIIIL 4.1, FIVL 2.9, TL 14.6, SHL 14.4, FOL 13.5, IMT 2.7, weight in life 2.4 g. Variation: Measurements of the type series are shown in Table 3. Paratypes and referred specimens generally agree with the holotype morphologically, but with the following exceptions: in VNMN 2017.003, AMS R177676 the middorsal ���U���-shaped ridge (on the holotype) is replaced by an inverted ���V���-shaped ridge. AMS R177676 with distinct dark grey blotches on ventral surface of thighs, this is lacking in VNMN 2017.003 and the holotype specimen. In life, AMS R177676 has a vivid red-orange flash on the axillary region (Fig. 4G). Gular region of VNMN 2017.002 and AMS R177678 lack dark blotches, present in all other specimens. Ventral surface of thighs (on AMS R177679 and AMS R177676) are covered in dark blotches, which are not present on any of the other specimens. Tongue weakly notched in one specimen (HNLP 2016062200001). Secondary sexual characters: Females of Megophrys rubrimera sp. nov. are currently unknown. Two male specimens (AMS R177678 and AMS R177679) possessed raised oval nuptial pads covered in microspinules on the base of Finger II; protruding fleshy projection (secondary sexual characteristic of some male Megophrys e.g. M. caudoprocta, M. koui and M. pachyproctus) above the cloaca absent from all specimens. After fixation it was very difficult to open the mouths of the specimens without damaging them due to the small head size. It was therefore not possible to determine the position, presence or absence of internal vocal slits which are a secondary sexual characteristic used as a diagnostic character to the species level (e.g. Mahony et al. 2013). Tadpole: (Fig. 5): The following tadpole description is based on a single specimen (AMS R177680) at Stage 37, the tadpole was confirmed as that of M. rubrimera sp. nov. by molecular analysis. Body elongated and slender; nares oval and are closer to the eye than to the snout; internarial distance is subequal to the interorbital distance; eyes positioned dorsolaterally, the pupils are round; the spiracle is sinistral and the spiracular tube protrudes from midway along the body wall just posterior to maximal trunk width and opens laterally; the tail makes up 68% of the total body length; the dorsal tail fin is low, particularly at the proximal half of the tail length; the basal tail width is 60% of the maximal trunk width; the oral disk is subterminal and antero-dorsal; the width of the umbelliform oral disc makes up ~80% of the maximal width of the trunk; the upper and lower lips each have deep medial emarginations; submarginal papillae are present on both sides between the oral orifice and the emarginations of the upper and lower lip, concentrated in a row around the margins; keratodonts are absent. Colour in life: Dorsally, the head and body are brown with darker speckles; obvious pale neuromasts; the oral disk is translucent yellowish brown with dark brown submarginal papillae; the dorsal and ventral fins opaque, pale yellowish brown with speckles; the venter is speckled white and brown; iris orange, speckled with black dots. Colour in preservative: Body brown with darker speckles; oral disc translucent grey brown with dark brown submarginal papillae; the dorsal and ventral fins opaque, pale grey brown with darker speckles; the venter is speckled grey and brown. Tadpole body measurements (in mm): BL 10.5, BH 4.4, BS 4.3, ES 1.7, IND 1.7, IOD 4.1, BW 5.4, ED 1.2, LFH 1.3, MTH 5.9, NE 1.9, ODW 4.5, SN 0.6, SS 5.7, TAL 22.8, TTL 33.3, TMH 3.1, TMW 5.4, UFH 1.3). Advertisement call: Call descriptions are based on the calls of one paratype (AMS R177677) and two unvouchered individuals. Advertisement calls were recorded at 21.0���22.9 ��C ambient temperature. Calls were an average of 73.3 ms (62���85 ms) in duration (Table 4; Fig. 6). Calls of unvouchered individual (b) contained an average of 23.05 pulses (19���25), whereas the calls of AMS R177677 and another unvouchered individual (a) were not distinctly pulsed. The average dominant frequency of calls was 3.3 kHz (3.2���3.4 kHz). Calls were repeated at a rate of approximately 3.25 (3.05���3.37) calls per second, and had an average intercall interval of 221.3 ms (190���261 ms). The number of calls within each call group ranged from 16���51, with an average of 38.7. Most call groups began at a relatively low amplitude, increasing with each call up to approximately a quarter to a third of the duration, after which amplitude remained relatively constant (Fig. 6A i ���ii & B i���ii). Individual calls either began with a medium relative amplitude and peaked near the middle before declining towards the end (Fig. 6A iii), or peaked at the beginning, steadily declining throughout (Fig. 6B iii). Natural history: All specimens of Megophrys rubrimera sp. nov. were found in disturbed evergreen secondary forest. Males were calling during June 2012 and 2016, and tadpoles (Stage 37) were collected in June 2016, suggesting a prolonged breeding season or that the species has a long larval period. Males were observed calling on stream-side vegetation (Fig. 7A���B) and a steep north-eastern facing roadside bank with seepages and a small stream (Fig. 7C���E). Females were not observed. Distribution and conservation status: This species is known from between 1400 m asl and 1722 m asl at two localities, 2 km apart in Sa Pa District, Lao Cai Province, Vietnam and 50 km north-west at Maandi, Jinping County, Ailao Mountain Range, Yunnan Province, China (Fig. 1; Chen et al. 2017). The species' area of occupancy (AOO) and extent of occurrence (EOO) are currently predicted to be 385 km 2 and 2298 km 2, respectively. The Vietnamese and Chinese portions of this species��� range comprise two threat-defined locations; land use likely differs between the two countries, and habitat loss probably affects the species independently in each. Ongoing disturbance to the species��� habitat due to forest clearance for agriculture has been observed in Sa Pa District. We recommend that Megophrys rubrimera sp. nov. is listed as Endangered in accordance with the IUCN Red List of Threatened Species categories and criteria B1ab(iii). Comparisons: Megophrys (Panophrys) rubrimera sp. nov. can be distinguished from all other congeners found in mainland southeast Asia, north of the Isthmus of Kra and neighbouring provinces of China on the basis of morphology, and from all congeners for which comparable data is available on the basis of molecular and acoustic data. Comparisons with each subgenus are discussed separately below. Subgenus Brachytarsophrys: Megophrys rubrimera sp. nov. can be distinguished from the species in the subgenus Brachytarsophrys (M. carinense, M. feae and M. intermedia) in mainland southeast Asia and neighbouring provinces of China by the absence of a transverse fold at the base of the head (versus presence), and having a smaller adult male size, SVL 26.7���30.5 mm (versus SVL> 79.1 mm; examined material). Subgenus Ophryophryne: Megophrys rubrimera sp. nov. can be distinguished from M. elfina by the presence of lateral fringes on the toes (versus absence; Poyarkov et al. 2017) and by the presence of vomerine teeth (versus absence Poyarkov et al. 2017); from M. gerti by having a smaller adult male size, SVL 26.7���30.5 mm (versus 31.7���42.2 mm; Poyarkov et al. 2017; material examined); from M. hansi by having a smaller adult male size, SVL 33.4���43.1 mm; Ohler 2003; Poyarkov et al. 2017; material examined); from M. microstoma by having a smaller adult male size, SVL 26.7���30.5 mm (versus 34.3���44.4 mm; Poyarkov et al. 2017; material examined; from M. koui by the absence of a protruding fleshy projection above the cloaca in sexually mature males (secondary sexual characteristic of sexually mature M. koui); from M. synoria by having a smaller adult male size, SVL 26.7���30.5 mm (versus 38.2���53.7 mm; Stuart et al. 2006b; Poyarkov et al. 2017; material examined) and further from all species in the subgenus Ophryophryne due to the presence (versus absence) of maxillary teeth. FFIGURE 7. Megophrys rubrimera sp. nov. in situ, and habitat at collection localities. (A) Adult male paratype AMS R177675 and (B) unvouchered calling male. (C���E) Habitat at type locality in Sa Pa District, Lao Cai Province, Vietnam. Subgenus Xenophrys: Megophrys rubrimera sp. nov. differs from M. auralensis by having a smaller adult male size, SVL 26.7���30.5 mm (versus 60.1���76.7 mm; Ohler et al. 2002; Neang et al. 2013), and a lack of interdigital toe webbing (versus rudimentary webbing); from M. damrei by having a smaller adult male size, SVL 26.7���30.5 mm (versus 47.5���57.1 mm: Mahony 2011; Neang et al. 2013; material examined), a lack of interdigital toe webbing (versus rudimentary), and the presence of lateral fringes on the toes (versus absence); from M. glandulosa by having a smaller adult male size, SVL 26.7���30.5 mm (versus 76.7���81.6 mm: Fei et al. 2009; material examined), and a lack of interdigital webbing between the toes (versus basal webbing); from M. lekaguli by having a smaller adult male size, SVL 26.7���30.5 mm (versus 55.6���68.1, Published as part of Tapley, Benjamin, Cutajar, Timothy, Mahony, Stephen, Nguyen, Chung Thanh, Dau, Vinh Quang, Nguyen, Tao Thien, Luong, Hao Van & Rowley, Jodi J. L., 2017, The Vietnamese population of Megophrys kuatunensis (Amphibia: Megophryidae) represents a new species of Asian horned frog from Vietnam and southern China, pp. 465-492 in Zootaxa 4344 (3) on pages 473-481, DOI: 10.11646/zootaxa.4344.3.3, http://zenodo.org/record/1043687, {"references":["Orlov, N. L., Khalikov, R. G., Murphy, R. w. & Lathrop, A. (2000) Atlas of Megophryids (Megophryidae: Anura: Amphibia) of Vietnam. [CD-ROM]","Orlov, N. L., Murphy, R. w., Ananjeva, N. B., Ryabov, S. A. & Ho, C. T. (2002) Herpetofauna of Vietnam, a checklist. Part 1. Amphibia. Russian Journal of Herpetology, 9 (2), 81 - 104.","Nguyen, V. S., Ho, T. C. & Nguyen, Q. T. (2009) Herpetofauna of Fietnam, Edition Chimaira, Frankfurt am Main, 768 pp.","Mahony, S., Teeling, E. C. & Biju, S. D. (2013) Three new species of horned frogs, Megophrys (Amphibia: Megophryidae), from northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region. Zootaxa, 3722 (2), 143 - 169. https: // doi. org / 10.11646 / zootaxa. 3722.2.2","Chen, J. M., Zhou, w. w., Poyarkov Jr, N. A., Stuart, B. L., Brown, R. M., Lathrop, A., wang, Y. Y., Yuan, Z. Y., Jiang, K., Hou, M. & Chen, H. M. (2017) A novel multilocus phylogenetic estimation reveals unrecognized diversity in Asian horned toads, genus Megophrys sensu lato (Anura: Megophryidae). Molecular Phylogenetics and Evolution, 106, 28 - 43. https: // doi. org / 10.1016 / j. ympev. 2016.09.004","Ohler, A. (2003) Revision of the genus Ophryophryne Boulenger, 1903 (Megophryidae) with description of two new species. Alytes, 1 - 2 (21), 23 - 44.","Stuart, B. L., Sok, K. & Neang, T. (2006 b) A collection of amphibians and reptiles from hilly eastern Cambodia. The Raffles Bulletin of Zoology, 54 (1), 129 - 155.","Ohler, A., Swan, S. R. & Daltry, J. C. (2002) A recent survey of the amphibian fauna of the Cardamom Mountains, southwest Cambodia with descriptions of three new species. The Raffles Bulletin of Zoology, 50 (2), 465 - 481.","Neang, T., Chhin, S., Meang, M. & Hun, S. (2013) Confirmation of three species of megophryid frogs (Amphibia Megophryidae) from the Cardamom Mountains of Southwest Cambodia, with the rediscovery of a long lost species. Cambodian Journal of Natural History, 2013 (2), 66 - 72.","Stuart, B. L., Chuaynkern, Y., Chan-ard, T. & Inger, R. F. (2006 a) Three new species of frogs and a new tadpole from eastern Thailand. Fieldiana Zoology, 111, 1 - 19. https: // doi. org / 10.3158 / 0015 - 0754 (2006) 187 [1: TNSOFA] 2.0. CO; 2","Bourret, R. (1937) Notes herpetologiques sur L'Indochine Francaise. XIV. Les batraciens de la collection du Laboratoire des Sciences Naturelles de l'Universite. Descriptions de quinze especes ou varietes nouvelles. Annexe au Bulletin General de l'Instruction Publique, 1937, 5 - 56.","Huang, Y. Z. & Fei, L. (1981) Two new amphibians from Tibet. Acta Zootaxonomica Sinica, 6, 211 - 215.","Li, Y. L., Jin, M. J., Zhao, J., Liu, Z. Y., wang, Y. Y. & Pang, H. (2014) Description of two new species of the genus Megophrys (Amphibia: Anura: Megophryidae) from Heishiding Nature Reserve, Fengkai, Guangdong, China, based on molecular and morphological data. Zootaxa, 3795 (4), 449 - 471. https: // doi. org / 10.11646 / zootaxa. 3795.4.5","Boulenger, G. A. (1899) On a collection of reptiles and batrachians made by Mr. J. D. La Touche in N. w. Fokien, China. Proceedings of the Zoological Society of London, 1899, 159 - 172, pls. XVI - XIX.","Rao, D. Q. & Yang, D. T. (1997) The karyotypes of Megophryinae (Pelobatidae) with a discussion on their classification and phylogenetic relationships. Asiatic Herpetological Research, 7, 93 - 102. https: // doi. org / 10.5962 / bhl. part. 18858","Fei, L., Ye, C. Y. & Jiang, J. P. (2012) Colored Atlas of Chinese Amphibians and their Distributions. Sichuan Publishing House of Science & Technology, Chengdu, 620 pp.","Orlov, N. L., Poyarkov Jr., N. A. & Nguyen, T. T. (2015) Taxonomic Notes on Megophrys Frogs (Megophryidae: Anura) of Vietnam, with Description of a New Species. Russian Journal of Herpetology, 22 (3), 206 - 18."]}

Megophrys (Panophrys) kuatunensis Pope, 1929 Figs. 8 & 9; Table 5. Megalophrys kuatunensis Pope 1929:1 (partim: see Remarks section). ������ Megophrys kuatunensis Gee & Boring 1929:20. ������ Megophrys (Megophrys) kuatunensis Dubois 1980:472. ������ Panophrys kuatunensis Rao and Yang 1997:98. ������ Xenophrys kuatunensis Delorme et al. 2006:17. ������ Megophrys (Panophrys) kuatunensis Mahony et al. 2017:755. Holotype: Adult male (AMNH 30126), type locality: ���Kuatun [Village], Chungan Hsien, northwest Fukien Province, China, 5500���6000 feet.��� (=Guadun [ca. 27��40���N, 117��40���E, ca. 1675���1830 m asl], Wuyishan County, Nanping Prefecture, Fujian Province), collected by Clifford H. Pope, April���September 1926 (Pope 1929, 1931). Paratypes: AMNH 30123���30124; AMNH 30239���30258, FMNH 24406 (formerly AMNH 30230), FMNH 24408 (formerly AMNH 30232), FMNH 24411���24413 (formerly AMNH 30235���30237), BMNH 1961.956 (formerly AMNH 30238, then FMNH 24414), BMNH 1985.1294 (formerly AMNH 30234, then FMNH 24410), BMNH 1985.1295 (formerly AMNH 30231, then FMNH 24407), MCZ 28297 (formerly AMNH 30233, then FMNH 24409). Examined specimens: Full morphological datasets were taken for five adult males, holotype and paratypes (AMNH 30126, 30240���30241, 30243���30244), and four adult female paratypes (AMNH 30242, BMNH 1961.956, BMNH 1985.1295, FMNH 24411). SVL measurements were taken for three additional adult male paratypes (FMNH 24406, FMNH 24412���24413). Holotype description: (Figs. 8 & 9; Table 5 for measurements): Sexually mature adult male. Head moderately small, width subequal to length; snout rounded in dorsal view, obtusely protruding in lateral view, without rostral appendage (Fig. 8); loreal region vertical and weakly concave; canthus rostralis angular; dorsal region of snout slightly concave; eye diameter greater than three times as long as maximum tympanum diameter, and longer than snout; eye-tympanum distance subequal to maximum tympanum diameter; tympanum circular, its upper edge not concealed by supratympanic ridge (Fig. 8); nostril orientated laterally, closer to eye than snout; internarial distance greater than eyelid width, and subequal to narrowest point between upper eyelids; pineal ocellus not visible externally; vomerine ridges and vomerine teeth absent; tongue moderately large, weakly notched posteriorly, with no medial lingual process. Forelimbs moderately short and thin, forearm moderately enlarged relative to upper forelimb, and shorter than hand; fingers short and narrow without lateral fringes (Fig. 9B), finger length formula IV Colouration in preservative: (Figs. 8 & 9): Dorsal and lateral surfaces of head, body, forelimbs and hindlimbs primarily mid-brown; solid dark brown triangular marking on dorsal surface of head between eyes; a dark brown ���X���-shaped marking on dorsum, with scattered dark brown spots and blotches on dorsum around ���X���- shaped marking and on dorsal surface of snout; front of snout, lateral canthus rostralis and lower half of supratympanic folds dark brown; wide vertical dark brown bar below eyes; dark brown blotch covering tympanum present; a short longitudinal dark brown stripe on central dorsum of snout; two dark brown crossbars on dorsal surface of forearms; dorsal surface of outer three fingers with dark brown transverse blotches; dorsal surface of hindlimbs with large dark brown transverse crossbars, two on thighs, one on shank, and one on tarsus, with dorsal surfaces of feet spotted. Ventral surfaces of head, body and limbs primarily light brown, with dark brown blotches along outer margin of ventral mandibles, gular region, chest and abdomen; a longitudinal dark brown blotch positioned medially, extending from the posterior gular region onto chest; a dark brown stripe extends laterally from the rear of the mandible, over the pectoral region and ventral proximal surface of forelimbs to approximately 70% distance to groin on both sides; ventral surfaces of thighs and shanks without markings; ventral surfaces of tarsus and feet dark brown; area surrounding vent and posterior surfaces of thighs dark brown; ventral surfaces of hands plain grey-brown and forearms ventrally with large dark brown blotch. Colouration in life: Not recorded. Variation: Refer to Table 5 for morphometric variation for five adult males and four adult females. Adult male SVL for eight paratypes ranges from 28.3 to 31.4 mm. Finger length formula varied slightly, IV ŠI ���), parietoscapular-middorsal ridge (���>������), complete parietoscapular-sacral ridge (���>- Secondary sexual characters: Males: weakly raised nuptial pads present, appearing smooth and translucent, or covered with brown microspinules, covering most of the dorsal surface of Finger I, narrowing distally and extending to the base of the distal phalange; nuptial pad absent on Finger II; vocal sac indistinct in preservation; internal vocal slits present near the rear of the mandible; forearms slightly to moderately enlarged relative to the upper forelimbs. Females: mature ova without pigmented poles; nuptial pads, internal vocal slits and enlarged forearms absent. Distribution (Fig. 10): To our knowledge, the presence of M. kuatunensis has only been positively confirmed with molecular data in Fujian and Jiangxi provinces (Li et al. 2014; Wang et al. 2014; Chen et al. 2017). The IUCN Red List assessment for this species indicates a much wider distribution (Huiqing et al. 2004) including locations in Zhejiang, Hunan and Guangxi provinces. Geological barriers exist between these locations that may limit the dispersal of Megophrys. The identity of populations assigned to M. kuatunensis in Zhejiang, Hunan and Guangxi provinces should be confirmed with molecular data. Remarks: Pope (1929) provided a relatively detailed description of the holotype of his new species, M. kuatunensis, including a brief comparison with M. boettgeri and M. minor. He later expanded on this description providing a figure of the holotype, a detailed variation section, and morphological and biological comparison with the sympatric M. boettgeri (Pope 1931). The type series of Megophrys kuatunensis was based on the holotype and 31 paratypes (Pope 1929). Pope (1931:445) included an additional specimen as a paratype, AMNH 30324, stating that ���An additional specimen has been found since the appearance of the original description which listed only 31 paratypes ���. This action does not validate AMNH 30324 as a paratype, and thus this specimen is not available for lectotype designation in the future should the holotype become lost/destroyed (International Commission of Zoological Nomenclature 1999). Within the section titled ���Notes on paratypes ���, Pope (1929) includes a statement that ���Tadpoles were secured���, neither designating a museum number, nor counting them among the 31 specimens referred to directly as paratypes, indicating that Pope did not intend to regard these tadpoles as paratypes. Pope (1931) further comments that of the two series of tadpoles (AMNH 30606 and AMNH 30645) collected from the type locality, he would only tentatively assign the former to M. kuatunensis. These tadpoles should not be considered to be paratypes of M. kuatunensis. Marx (1958) lists three paratypes in the CNHM (FMNH 24408, 24412, 24413), omitting two paratypes FMNH 24406 (formerly AMNH 30230) and FMNH 24411 (formerly AMNH 30235) from his annotated catalogue of type specimens in the collection. This omission was not amended in supplementary catalogue of FMNH types (Marx 1976), however, we confirm that both of these specimens were present in the collection during a recent visit (S. Mahony, pers. com.). In their description of a new species, M. brachykolos, Inger and Romer (1961) examined part of the type series of M. kuatunensis and were the first to note that the paratype series of M. kuatunensis is heterogeneous in species composition. They identified a M. kuatunensis female paratype FMNH (as CNHM) 24408 as M. brachykolos on the basis of several measurements and characters that fell within the variation range of the female M. brachykolos types. They designated this specimen as a paratype of their new species (Inger and Romer 1961; Marx 1976). Our examination of approximately half of the type series of M. kuatunensis (including FMNH 24408) revealed two additional paratypes (AMNH 30239 and BMNH 1985.1294 [formerly AMNH 30234, then FMNH 24410]) that are morphologically conspecific with FMNH 24408, and not M. kuatunensis sensu stricto. Though we partially agree with Inger and Romer (1961) in so far as these specimens share superficial similarities with M. brachykolos sensu stricto (and its sister taxon M. acuta), we suggest that their identification should be considered tentative pending further study of this population due to geographic distance from the type locality of M. brachykolos, and the recognition of extensive cryptic diversity in Megophrys (Chen et al. 2017; Mahony et al. 2017)., Published as part of Tapley, Benjamin, Cutajar, Timothy, Mahony, Stephen, Nguyen, Chung Thanh, Dau, Vinh Quang, Nguyen, Tao Thien, Luong, Hao Van & Rowley, Jodi J. L., 2017, The Vietnamese population of Megophrys kuatunensis (Amphibia: Megophryidae) represents a new species of Asian horned frog from Vietnam and southern China, pp. 465-492 in Zootaxa 4344 (3) on pages 482-486, DOI: 10.11646/zootaxa.4344.3.3, http://zenodo.org/record/1043687, {"references":["Pope, C. H. (1929) Four new frogs from Fukien, China. American Museum Novitates, 352, 1 - 5.","Gee, N. G. & Boring, A. M. (1929 [\" 1930 \"]) A check list of Chinese Amphibia with notes on geographical distribution. Peking Society of Natural History Bulletin, 4 (2), 15 - 51.","Rao, D. Q. & Yang, D. T. (1997) The karyotypes of Megophryinae (Pelobatidae) with a discussion on their classification and phylogenetic relationships. Asiatic Herpetological Research, 7, 93 - 102. https: // doi. org / 10.5962 / bhl. part. 18858","Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anoures). Alytes, 24 (1 - 4), 6 - 21.","Mahony, S., Foley, N., Biju, S. D. & Teeling, E. (2017) Evolutionary History of the Asian Horned Frogs (Megophryinae): Integrative Approaches to Timetree Dating in the Absence of a Fossil Record. Molecular Biology, 34 (3), 744 - 771. https: // doi. org / 10.1093 / molbev / msw 267","Pope, C. H. (1931) Notes on amphibians from Fukien, Hainan, and other parts of China. Bulletin of the AMNH, 61 (Article 8), 1 - 215.","Li, Y. L., Jin, M. J., Zhao, J., Liu, Z. Y., wang, Y. Y. & Pang, H. (2014) Description of two new species of the genus Megophrys (Amphibia: Anura: Megophryidae) from Heishiding Nature Reserve, Fengkai, Guangdong, China, based on molecular and morphological data. Zootaxa, 3795 (4), 449 - 471. https: // doi. org / 10.11646 / zootaxa. 3795.4.5","Chen, J. M., Zhou, w. w., Poyarkov Jr, N. A., Stuart, B. L., Brown, R. M., Lathrop, A., wang, Y. Y., Yuan, Z. Y., Jiang, K., Hou, M. & Chen, H. M. (2017) A novel multilocus phylogenetic estimation reveals unrecognized diversity in Asian horned toads, genus Megophrys sensu lato (Anura: Megophryidae). Molecular Phylogenetics and Evolution, 106, 28 - 43. https: // doi. org / 10.1016 / j. ympev. 2016.09.004","Huiqing, G., Baorong, G., van Dijk, P. P. & Ohler, A. (2004) Megophrys kuatunensis. The IUCN Red List of Threatened Species 2004: e. T 57641 A 11668487. https: // doi. org / 10.2305 / IUCN. UK. 2004. RLTS. T 57641 A 11668487. en","Marx, H. (1976) Supplementary catalogue of type specimens of reptiles and amphibians in Field Museum of Natural History. Field Museum of Natural History, Chicago. [unknown pagination]","Inger, R. F. & Romer, J. D. (1961) A new Pelobatid frog of the genus Megophrys from Hong Kong. Chicago Natural History Museum, 39 (46), 533 - 538."]}