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Calamaria dominici Ziegler, Tran & Nguyen sp. nov. Figs1-4 Holotype: IEBR A.2018.1, an adult female collected on 28 May 2017 at 11:30 on a forest path by Anh Vu Tran in evergreen mixed forest of broadleaf and conifer trees within Ta Dung Nature Reserve, Dak Nong Province, Central Highlands, Vietnam, at an elevation of 1240 m asl. Diagnosis: A species of the genus Calamaria cha- racterized by the combination of the following characters: (1) rostral wider than high; (2) paraparietal surrounded by six shields and scales; (3) eye diameter larger than eye-mouth distance; (4) preocular present; (5) supralabials 5/4, 3-4/2-3 entering orbit; (6) maxillary teeth nine, modified; (7) infralabials 5/4, first three touching anterior chin shields; (8) mental touching tip of right anterior chin shield; (9) ventrals 1 + 174; subcaudal scales 18/17, divided; (10) precloacal plate single; (11) tail relatively short (6.2% of the total length), nearly as thick as body, slightly tapering, and ending in obtuse point; (12) dorsal scales reducing to six rows at position above 4th subcaudal, and to four rows above 13th subcaudal on tail; (13) dorsum dark with irregular yellow blotches; and (14) ventral side dark with few yellow blotches and bands. Description of holotype: Habitus vermiform; head indistinct from neck; pupil round; tail relatively short (6.2% of the total length), nearly as thick as body, slightly tapering, ending in obtuse point. Size. SVL: 395 mm; TaL: 26 mm; TL: 421 mm; ratio TaL/TL: 0.06. Dentition. Right upper maxilla with 9 modified maxillary teeth. Body scalation. Dorsal scale rows 13���13���13, smooth. Dorsal scales reducing to six rows at position above fourth subcaudal, to five rows above 12th subcaudal, and to four rows above 13th subcaudal on tail. 174 ventrals (+ 1 preventral); 18/17 subcaudals, all paired, first pair not in contact, followed by tail tip; precloacal single. Head scalation. Rostral wider than high, portion visible from above shorter than prefrontal suture. Prefrontal shorter than frontal, not entering orbit, and touching first two supralabials on right side and second and third supralabial on left side. Frontal hexagonal, nearly two times maximum width of supraocular. Paraparietal surrounded by six shields and scales. Length of parietal shorter than distance from posterior tip of frontal to posterior tip of rostral. Supraocular 1/1. Loreal 1/1. Preocular 1/1, distinctly higher than wide. Postocular 1/1, higher than wide, not as high as eye diameter. Eye diameter larger than eye-mouth distance. 5/4 supralabials, on left side third and fourth entering orbit, fifth longest; on right side second and third entering orbit, fourth longest. Mental triangular, touching tip of right anterior chin shields. 5/4 infralabials, first three touching anterior chin shields. First pair of chin shields in contact mesially, second pair touching anteriorly and separated posteriorly. Coloration (in life). Eye black; tongue grey anteriorly, pinkish-grey posteriorly; dorsum of body and tail dark purplish-black, iridescent; head with irregular small and few medium-sized yellow blotches; body with mediumsized to large yellow blotches, irregularly arranged, in part forming transverse or oblique rows, sometimes in zig-zag pattern; hind part of body dorsum and dorsal surface of tail less intensely blotched; venter purplishblack with irregular yellow transverse bands or blotches; chin and throat region dark with yellow reticulation or blotches and bands; lower tail surface dark with few yellow blotches or bands. Coloration (in preservative). Ground colour purplishblack to brownish-black with whitish-cream pattern of irregularly arranged blotches. Comparisons: Comparisons of the new species with its congeners took place based on the following references (Inger & Marx, 1965; Grismer et al., 2004, Howard & Gillespie, 2007, Ziegler et al., 2008; Koch et al., 2009; Nguyen et al., 2009; Orlov, 2009; Orlov et al., 2010). In the following we first compare Calamaria dominici sp. nov. with the Calamaria species reported to occur in Vietnam: Calamaria dominici sp. nov. already differs by its color pattern from the species known from Vietnam: C. abramovi has a black dorsum without spots, the venter is covered with yellow-orange spots; C. buchi is blackish above with each dorsal scale having small light spots and its ventral scales having dark outermost corners; C. concolor has a uniform, patternless light brown body dorsum and a cream venter; C. gialaiensis has a light greyish brown dorsum with an indistinct dark neck collar and few dark blotches along posterior vertebral region, two pairs of light blotches on the tail, as well as a yellowish beige ventral side, with dark outermost corners of ventrals and anterior subcaudals; C. pavimentata usually has narrow, dark, longitudinal stripes, and a solid black collar behind the neck; C. sangi has a greyish brown dorsum with fine dark mottling, as well as a yellowish beige ventral side, with dark transverse bands and a dark longitudinal stripe below the tail; C. septentrionalis has dorsal scales with many small light dots forming a network; C. lovii ingermarxorum has an immaculate grey-bluish dorsum with light spots on each side of the neck covering four scales; and C. thanhi has distinct transverse light body bands. Calamaria dominici sp. nov. further differs from the species so far known from Vietnam in morphological characters: Calamaria dominici sp. nov. differs from C. abramovi Orlov, 2009 by its rostral being wider than high (versus its height equal to width), by the dorsal scales reducing to four rows above 13th subcaudal on tail (versus above 20th subcaudal in the female holotype of C. abramovi), by 18/17 versus 20 subcaudals in females, and by having 13-13-13 versus 12-13-13 dorsal scale rows. Calamaria dominici sp. nov. differs from C. buchi Marx & Inger, 1955 by having fewer ventral scales in females (1 + 174 versus 221-236 in C. buchi), by the rostral being wider than high (versus rostral higher than wide), by the dorsal scales reducing to four rows above 13th subcaudal on tail (versus above 3rd-4th subcaudal), by 18/17 versus 13-14 subcaudals in females, by having a ratio of tail length to total length of 6.2% (versus 3.9-4.1 in female C. buchi), and by the length of parietal being shorter than distance from posterior tip of frontal to posterior tip of rostral (versus length of parietal greater than distance from posterior tip of frontal to posterior tip of rostral). Calamaria dominici sp. nov. differs from C. concolor Orlov, Nguyen, Nguyen, Ananjeva & Ho, 2010 by having paraparietal surrounded by six shields and scales (versus by five shields and scales), and by the dorsal scales reducing to four rows above 13th subcaudal on tail (versus above 19th subcaudal in the male holotype of C. concolor). Calamaria dominici sp. nov. differs from C. gialaiensis Ziegler, Nguyen & Nguyen, 2009 by having paraparietal surrounded by six shields and scales (versus by five shields and scales), and by tail ending in obtuse point (versus with rounded end). Calamaria dominici sp. nov. differs from C. lovii Boulenger, 1887 by having a preocular scale (versus being absent in C. lovii), and by tail ending in obtuse point (versus with blunt end in the subspecies occurring in Vietnam, C. lovii ingermarxorum). Calamaria dominici sp. nov. differs from C. pavimentata Dum��ril, Bibron & Dum��ril, 1854 by the rostral being wider than high (versus rostral as broad as high or slightly higher than broad), and by the tail ending in obtuse point (tail tip with sharp point in C. pavimentata). Calamaria dominici sp. nov. differs from C. sangi Nguyen, Koch & Ziegler, 2009 by having fewer ventral scales (1 + 174 versus 2 + 190 in C. sangi), and by the dorsal scales reducing to six rows above 4th subcaudal on tail (versus above 8th subcaudal). Calamaria dominici sp. nov. differs from C. septentrionalis Boulenger 1890 by 18/17 versus 6-11 subcaudals in females, by having a ratio of tail length to total length of 6.2% (versus 2.6-4.3 in female C. septentrionalis), and by the tail ending in obtuse point (versus tail tip rounded in C. septentrionalis). Calamaria dominici sp. nov. differs from C. thanhi Ziegler & Le, 2005 by having a preocular scale (versus absent), by having fewer ventral scales (1 + 174 versus 198 in C. thanhi), and by the tail ending in obtuse point (tail tip with sharp point in C. thanhi). Calamaria dominici sp. nov. differs from C. yunnanensis Chernov, 1962, a species reported from southern China, which was judged as a doubtful form by Inger & Marx (1965), but subsequently listed as valid by Yang & Inger (1986) and Zhao & Adler (1993), by having a preocular (versus being absent), and by lacking narrow, dark, elongated stripes along the body. Calamaria dominici sp. nov. has a distinct preocular scale, which is lacking in the following species: C. alidae Boulenger, 1920, C. apraeocularis Smith, 1927, C. banggaiensis Koch, Arida, McGuire, Iskandar & B��hme, 2009, C. butonensis Howard & Gillespie, 2007, C. ceramensis De Rooij, 1913, C. gracillima (G��nther, 1872), C. javanica Boulenger, 1891, C. longirostris Howard & Gillespie, 2007, C. mecheli Schenkel, 1901, C. rebentischi Bleeker, 1860, and C. schmidti Marx & Inger, 1955. Calamaria dominici sp. nov. has modified maxillary teeth and thus differs from the following species, which have unmodified maxillary teeth: C. acutirostris Boulenger, 1896, C. curta Boulenger, 1896, C. lautensis De Rooij, 1917, C. leucogaster Bleeker, 1860, and C. ulmeri Sackett, 1940. Calamaria dominici sp. nov. has paraparietal surrounded by six shields and scales and thus differs from C. albiventer (Gray, 1835) (5), C. bicolor Dum��ril, Bibron & Dum��ril, 1854 (5), C. bitorques Peters, 1872 (5), C. brongersmai Inger & Marx, 1965 (5), C. everetti Boulenger, 1893 (5), C. griswoldi Loveridge, 1938 (5), C. hilleniusi Inger & Marx, 1965 (5), C. joloensis Taylor, 1922 (5), C. lateralis Mocquard, 1890 (5), C. lumbricoidea Boie, 1827 (4 or 5), C. lumholtzi Andersson, 1923 (5), C. muelleri Boulenger, 1896 (5), C. palavanensis Inger & Marx, 1965 (5), C. prakkei Lidth de Jeude, 1893 (5), and C. suluensis Taylor, 1922 (5). Calamaria dominici sp. nov. differs from the following species by a distinctly higher ventral scale count in the female sex: C. abstrusa Inger & Marx, 1965 (145-152), C. crassa Lidth de Jeude, 1922 (158-164), C. eiselti Inger & Marx, 1965 (151-153), C. linnaei Boie, 1827 (148- 166), and C. melanota Jan, 1862 (131-154). Calamaria dominici sp. nov. differs from the following species by a distinctly higher subcaudal scale count in the female sex: C. margaritophora Bleeker, 1860 (8- 11), C. nuchalis Boulenger, 1896 (9), and C. sumatrana Edeling, 1870 (10-14). Calamaria dominici sp. nov. differs from C. grabowskyi Fischer, 1885 by a distinctly lower subcaudal scale count in the female sex (20-28). Calamaria dominici sp. nov. has the first three infralabials touching the anterior chin shields versus only two pairs of infralabials touching anterior chin shields in C. borneensis Bleeker, 1860. In addition, Calamaria dominici sp. nov. differs from the remaining species at least by a distinct colour pattern: C. battersbyi Inger & Marx, 1965 (with narrow longitudinal stripes mid-dorsally), C. oesemani Inger & Marx, 1965 (with a continuous light stripe the entire length of the body), C. doederleini Gough, 1902 (with narrow dark brown crossbands on body and tail), C. forcarti Inger & Marx, 1965 (with narrow dark crossbands behind head, body without stripes, venter yellow), C. gervaisii Dum��ril, Bibron & Dum��ril, 1854 (usually with a dark-edged, interrupted, light stripe on first body scale row), C. ingeri Grismer, Kaiser & Yaakob, 2004 (with incomplete light transverse bands on body and tail), C. modesta Dum��ril, Bibron & Dum��ril, 1854 (with ventrals having dark pigment at least laterally), C. schlegeli Dum��ril, Bibron & Dum��ril, 1854 (dark above, light below, head yellow above and below or black above and yellow below, or intermediate conditions), C. virgulata Boie, 1827 (dorsally dark brown, each scale with a light network, with or without longitudinal dark stripes). Etymology: Named dominici to honor Dominic T. Charles Scriven, founder of Wildlife at Risk (WAR), for his contribution towards wildlife conservation in Vietnam. Suggested common names: Dominic���s reed snake (English), Ran mai gam do-mi-nic (Vietnamese), Calamaire de Dominic (French), and Dominics Zwergschlange (German). Distribution: Calamaria dominici is currently known only from the type locality (Fig. 5). Natural history: The holotype was found in evergreen mixed forest of broadleaf and conifer trees. The snake was discovered on a forest path near a small creek, for about 50 m distance from a large creek (Figs 6-7). It was found, surface active, in a densely vegetated boggy area at 11:30. The surrounding habitat was primary forest consisting of dense understory punctuated with large boulders scattered over a ca. 20�� slope that descended to a large creek. Dissection of the female holotype revealed ovaries with some eggs enlarge to 4 mm., Published as part of Ziegler, Thomas, Tran, Vu A., Babb, Randall D., Jones, Thomas R., Moler, Paul E., Van Devender, Robert W. & Nguyen, Truong Q., 2019, A new species of reed snake, Calamaria Boie, 1827 from the Central Highlands of Vietnam (Squamata: Colubridae), pp. 17-26 in Revue suisse de Zoologie 126 (1) on pages 18-23, DOI: 10.5281/zenodo.2619512, {"references":["Inger R. F., Marx H. 1965. The systematics and evolution of the oriental colubrid snakes of the genus Calamaria. Fieldiana: Zoology 49: 1 - 304.","Grismer L. L., Kaiser H., Yaakob N. S. 2004. A new species of reed snake of the genus Calamaria H. Boie, 1827, from Pulau Tioman, Pahang, West Malaysia. Hamadryad 28 (1 & 2): 1 - 6.","Howard S. D., Gillespie G. R. 2007. Two New Calamaria (Serpentes) Species from Sulawesi, Indonesia. Journal of Herpetology 41 (2): 237 - 242.","Ziegler T., Nguyen S. V., Nguyen T. Q. 2008. A new reed snake of the genus Calamaria Boie (Squamata: Colubridae) from Vietnam. Current Herpetology 27 (2): 71 - 80.","Koch A., Arida E., McGuire J. A., Iskandar D. T., Bohme W. 2009. A new species of Calamaria (Squamata: Colubridae) similar to C. ceramensis de Rooij, 1913, from the Banggai Islands, east of Sulawesi, Indonesia. Zootaxa 2196: 19 - 30","Nguyen T. Q., Koch A., Ziegler T. 2009. A new species of reed snake, Calamaria Boie, 1827 (Squamata: Colubridae), from central Vietnam. Hamadryad 34 (1): 1 - 8.","Orlov N. L. 2009. A new species of the genus Calamaria (Squamata: Ophidia: Colubridae) from the central highlands (Ngoc Linh Nature Reserve, Ngoc Linh Mountain, Kon Tum Province), Vietnam. Russian Journal of Herpetology 16 (2): 146 - 154.","Orlov N. L., Nguyen T. Q., Nguyen T. T., Ananjeva N. B., Ho C. T. 2010. A new species of the genus Calamaria (Squamata: Ophidia: Colubridae) from Thua Thien-Hue Province, Vietnam. Russian Journal of Herpetology 17 (3): 236 - 242.","Ziegler T., Le Q. K. 2005. A new species of reed snake, Calamaria (Squamata: Colubridae), from the Central Truong Son (Annamite mountain range), Vietnam. Zootaxa 1042: 27 - 38.","Yang D., Inger R. F. 1986. Key to the snakes and lizards of China. Smithsonian Herpetological Information Service 71: 1 - 21.","Zhao E., Adler K. 1993. Herpetology of China. Society for the Study of Amphibians and Reptiles, Contribution to Herpetology 10: 1 - 522."]}

Ziegler, Thomas, Tran, Vu A., Babb, Randall D., Jones, Thomas R., Moler, Paul E., Van Devender, Robert W., Nguyen, Truong Q. (2019): A new species of reed snake, Calamaria Boie, 1827 from the Central Highlands of Vietnam (Squamata: Colubridae). Revue suisse de Zoologie 126 (1): 17-26, DOI: 10.5281/zenodo.2619512, {"references":["Chen J.M., Poyarkov N.A., Suwannapoom C., Lathrop A., Wu Y.H., Zhou W.W., Yuan Z.Y., Jin J.Q., Chen H.M., Liu H.Q., Nguyen T.Q., Nguyen S.N., Duong T.V., Eto K., Nishikawa K., Matsui M., Orlov N.L., Stuart B.L., Brown R.M., Rowley J.J.L., Murphy R.W., Wang Y.Y, Che J. 2018. Large-scale phylogenetic analyses provide insights into unrecognized diversity and historical biogeography of Asian leaf-litter frogs, genus Leptolalax (Anura: Megophryidae). Molecular Phylogenetics and Evolution 124: 162-171.","Do T.V., Luu T.H., Wanke S., Neinhuis C. 2015. Three New Species and Three New Records of Aristolochia Subgenus Siphisia from Vietnam including a Key to the Asian Species. Systematic Botany 40(3): 671-691.","Dowling H.G. 1951. A proposed standard system of counting ventrals in snakes. British Journal of Herpetology, 1(1): 97-99.","Grismer L.L., Kaiser H., Yaakob N.S. 2004. A new species of reed snake of the genus Calamaria H. Boie, 1827, from Pulau Tioman, Pahang, West Malaysia. Hamadryad 28 (1&2): 1-6.","Howard S.D., Gillespie G.R. 2007. Two New Calamaria (Serpentes) Species from Sulawesi, Indonesia. Journal of Herpetology 41(2): 237-242.","Inger R.F., Marx H. 1965. The systematics and evolution of the oriental colubrid snakes of the genus Calamaria. Fieldiana: Zoology 49: 1-304.","Koch A., Arida E., McGuire J.A., Iskandar D.T., Bohme W. 2009. A new species of Calamaria (Squamata: Colubridae) similar to C. ceramensis de Rooij, 1913, from the Banggai Islands, east of Sulawesi, Indonesia. Zootaxa 2196: 19-30","Nguyen T.Q., Koch A., Ziegler T. 2009. A new species of reed snake, Calamaria Boie, 1827 (Squamata: Colubridae), from central Vietnam. Hamadryad 34(1): 1-8.","Orlov N. L. 2009. A new species of the genus Calamaria (Squamata: Ophidia: Colubridae) from the central highlands (Ngoc Linh Nature Reserve, Ngoc Linh Mountain, Kon Tum Province), Vietnam. Russian Journal of Herpetology 16(2): 146-154.","Orlov N.L., Nguyen T.Q., Nguyen T.T., Ananjeva N.B., Ho C.T. 2010. A new species of the genus Calamaria (Squamata: Ophidia: Colubridae) from Thua Thien-Hue Province, Vietnam. Russian Journal of Herpetology 17(3): 236-242.","Rowley J.J.L., Tran D.T A., Le D.T.T., Dau V.Q., Peloso P.L.V., Nguyen T.Q., Hoang H.D., Nguyen T.T., Ziegler T. 2016. Five new, microendemic Asian Leaf-litter frogs (Leptolalax) from the southern Annamite mountains, Vietnam. Zootaxa 4085(1): 63-102.","Simmons J.E. 2002. Herpetological collecting and collections management. Revised edition. Society for the Study of Amphibians and Reptiles. Herpetological Circular, 31: 1-153.","Uetz P., Freed P., Hosek J. 2018. Eds. The Reptile Database, http://www.reptile-database.org, accessed 20.06.2018.","Wallach V.K., Williams L., Boundy J. 2014. Snakes of the World: A Catalogue of Living and Extinct Species. Taylor and Francis, CRC Press, 1237 pp.","Yang D., Inger R.F. 1986. Key to the snakes and lizards of China. Smithsonian Herpetological Information Service 71: 1-21.","Zhao E., Adler K. 1993. Herpetology of China. Society for the Study of Amphibians and Reptiles, Contribution to Herpetology 10: 1-522.","Ziegler T., Le Q. K. 2005. A new species of reed snake, Calamaria (Squamata: Colubridae), from the Central Truong Son (Annamite mountain range), Vietnam. Zootaxa 1042: 27-38.","Ziegler T., Nguyen S.V., Nguyen T.Q. 2008. A new reed snake of the genus Calamaria Boie (Squamata: Colubridae) from Vietnam. Current Herpetology 27(2): 71-80."]}

Opisthotropis voquyi sp. nov. Holotype. IEBR 4326 [Field No. TYT.2013.4], an adult male, from Tay Yen Tu Nature Reserve, Bac Giang Province, 437 m asl., Vietnam, collected by Cuong The Pham et al. on 14 June 2013 (Figs. 6���8). Paratypes. VNMN 0 6315 [Field No. TYT 2013.1], male, collected by Cuong The Pham et al. in June 2013; VNMN 0 6316 [Field No. BG 2015.13], female, collected by Cuong The Pham et al. on 17 May 2015; IEBR 4327 [Field No. BG 2015.33], female, collected by Cuong The Pham et al. on 22 May 2015; IEBR 4328 [Field No. BG 2015.34], female, collected by Cuong The Pham et al. on 22 May 2015; ZFMK 100819 [Field No. BG 2015.35], female, collected by Cuong The Pham et al. on 22 May 2015; ZFMK 100820 [Field No. TYT 2014.11], female, collected by Hang An Thi in May 2014, all paratypes same locality data as the holotype (Figs. 8���10). Bourret & Pa) Sa Bourret & Pa) Sa Chapa () 1934 a? Tamdao) b Chapa) (a 1935 Chapa (a 1935) Chapa a (1935) Tam Dao (1935) b Dao Tam (b) 1935 NganSon c1935 ()) (1936 Angel 1933 (syntype Angel (1933 syntype. 259 M Bourret A. 57 1934 (. M 518 Bourret 953 X. Bourret. 954 X Bourret 451 M. Bourret 794. X Bourret M. 498 Bourret Bourret ...Continued on next page TABLE 2b. (Continued) Diagnosis. A species of the genus Opisthotropis, characterized by a combination of the following characters: (1) internasal not in contact with loreal; (2) one preocular; (3) usually two postoculars; (4) one anterior temporal; (5) one posterior temporal; (6) 7 or 8, rarely 9 supralabials; (7) 25 maxillary teeth; (8) subcaudals 74���86; (9) 15 dorsal scale rows at neck, at midbody and before vent; (10) body scales smooth or only with few faint keels; (11) dorsal scales being grey, greyish-brown or brown in preservative, posteriorly more or less edged with pale greyishbrown. Description of holotype. Body stout, oval in cross section; head short, slightly broader than neck, depressed, dorsally covered with large shields; nostrils in dorsal position and oblique, narrow, piercing the middle of nasal, which is divided beneath; eye small, pupil round. Size. SVL: 337 mm; TaL: 106 mm; TL: 443 mm; ratio TaL/TL: 0.239. Dentition. Maxillary teeth: right upper maxilla with 25 subequal teeth or sockets, without diastema. Body scalation. Dorsal scale rows 15���15���15, smooth, with few faint keels. 181 ventrals (+ 1 preventral); 85 subcaudals, all paired; precloacal divided. Head scalation. Rostral heptagonal, wider than high, readily visible from above; nasals large, much longer than high, divided below nostril by a distinct furrow; internasals two, strongly curved, longer than wide, in contact with rostral, nasals, and prefrontal; prefrontal single, anteriorly pointed, much broader than long, in contact with internasals, nasals, loreals, preoculars, and frontal; frontal pentagonal, slightly wider than long, apex directed posteriorly, 1.3 times longer than prefrontal; parietals longer than wide, in contact approximately the length of frontal; 1 / 1 supraocular, distinctly wider than high; 1 / 1 loreal, pentagonal, slightly wider than high, not in contact with internasal; 1 / 1 preocular, large, hexagonal, distinctly higher than wide, reaching frontal, in broad contact with prefrontal; 2 / 2 postoculars, upper largest, curved, higher than wide, lower distinctly wider than high; 8 / 7 supralabials, anterior ones distinctly higher than long, fourth entering orbit (the fourth supralabial on the right side fused with the subsequent supralabial), SL 7 / 6 largest; 1+1 / 1+1 temporals, anterior one very long and narrow, in broad contact with SL 6���7 / 5���6 and parietals, posterior one more strongly developed; infralabials 9 / 9, first pair in contact behind small mental, IL 1���4 in contact with anterior chin shields, IL 5 largest, apex directed posteriorly; posterior chin shields about the same length than anterior ones, separated from each other by 2 + 4 scales. Hemipenes. Only the left hemipenis fully everted, ending in two short, inwards curved lobes (Fig. 7). Sulcus spermaticus undivided, extending only to the slightly smaller lobe. Hemipenes covered with small spines, curved backwards. Only the terminal area of the hemipenes between the lobes without spines. Upper truncus of hemipenes with ring of enlarged spines. Largest spines extending from sulcus spermaticus to hemipenis sides, with the lowermost one being the most distinct. Lower truncus of hemipenes covered with tiny spines. Coloration (in preservative). Upper head and body surface brown, the venter yellowish-beige. Dorsal scales posteriorly more or less edged with pale brown. Dorsal head surface paler around nostrils and in the middle of parietals. Infralabials same colour as dorsal scales, adjoining scales becoming gradually paler towards mental groove. Outermost edges of ventrals and subcaudals the same colour as dorsal scales. Subcaudals speckled with small dark spots, becoming more numerous towards the tail tip. Variation of paratypes. An overview of the meristic and scalation data of the type series of Opisthotropis voqyui sp. nov. is given in Table 3. The dorsal ground coloration varies between greyish, greyish brown, and brown. Concerning a potential sexual dimorphism, males seem to be brown, females grey or greyish brown. Females also seem to grow larger. The dorsal tail scales in the males had few faint keels, which was only the case for one female; the remaining four females had smooth dorsal tail scales. The paratype series generally accorded well with the holotype description. Of the type series six individuals had two postoculars, only one individual had 2 postoculars on the left side and one postocular on the right side. Comparisons. The comparison of Opisthotropis voquyi sp. nov. with other species of the genus Opisthotropis and the genera Parahelicops and Paratapinophis was based on the following references: Stuart & Chuaynkern (2007), David et al. (2011), Teyni�� et al. (2013), Nguyen et al. (in press), and Wang et al. (2017). There are only few species of Opisthotropis and other genera with 15 midbody dorsal scale rows (see Table 4). 15 15 or 17 17 19 23 Opisthotropis alcalai Brown & Leviton, 1961 x O. andersonii (Boulenger, 1888) x O. atra G��nther, 1872 x O. balteata (Cope, 1895) x O. cheni Zhao, 1999 x O. cucae David, Pham, Nguyen & Ziegler, 2011 x O. daovantieni Orlov, Darevsky & Murphy, 1998 x O. durandi Teyni��, Lottier, David, Nguyen & Vogel, 2013 x O. guangxiensis Zhao, Jiang & Huang, 1978 x O. jacobi Angel & Bourret, 1933 x O. kikuzatoi Okada & Takara, 1958 x O. kuatunensis Pope, 1928 x O. lateralis Boulenger, 1903 x O. latouchii (Boulenger, 1899) x O. laui Yang, Sung & Chan, 2013 x ...Continued on next page Main additional differences between Opisthotropis voquyi sp. nov. and Parahelicops annamensis are: one preocular in the new species versus two in P. annamensis; internasal not in contact with loreal in the new species versus in contact in P. annamensis; body scales smooth or only with few faint keels in the new species versus keeled throughout in P. annamensis and especially strongly keeled on the posterior part of the body and the base of the tail; body dorsum grey to greyish-brown or brown, with dorsals posteriorly more or less edged with pale greyish-brown in the new species versus dorsum with orange bars and spots in P. annamensis. Opisthotropis voquyi sp. nov. differs from O. guangxiensis by 7���9 versus 9���10 supralabials, one posterior temporal (versus two), 74���86 versus 51���58 subcaudals, dorsal scales arranged in 15���15���15 versus 17���15���15 rows, and a greyish to greyish���brown or brown dorsum, dorsal scales posteriorly more or less edged with pale greyishbrown in the new species versus dark body with pale crossbars in O. guangxiensis. Opisthotropis voquyi sp. nov. differs from O. kikuzatoi by 7���9 supralabials (versus 6), one preocular (versus two), one anterior temporal (versus two), and the greyish to greyish-brown or brown dorsal pattern, with the dorsals posteriorly being more or less edged with pale greyish-brown in the new species (versus dark dorsum with dorsolateral orange spots). Opisthotropis voquyi sp. nov. differs from O. maculosa by a grey to greyish-brown or brown dorsum, with dorsals being posteriorly more or less edged with pale greyish-brown in the new species versus with yellow dots on a dark background in O. maculosa. In external morphological characters, O. voquyi sp. nov. is particularly similar to O. jacobi, from which it differs in usually having two postoculars versus only one in O. jacobi, in the number of maxillary teeth: 25 in the new species versus 19���23 in O. jacobi, and in the dorsal scales being grey to greyish���brown or brown versus blackish���brown in O. jacobi. Etymology. The specific epithet is a noun in the genitive case. This species is named in honour of Professor Doctor Vo Quy of Vietnam National University, Hanoi, who passed away on 10 January 2017, and who played a pioneering role in nature conservation in Vietnam. He inspired later generations of conservation biologists, including authors of this study. Suggested common names. Vo Quy���s Mountain Stream Keelback (English) and R���n tr��n v�� qu�� (Vietnamese). Distribution. Opisthotropis voquyi is currently known only from the type locality in Bac Giang Province (Fig. 11). Natural history. Specimens were found at night between 19:30 and 23:00 h in and around small rocky streams, where the snakes were observed in the water or on the rocks close to the water. The surrounding habitat was secondary evergreen forest of medium and small hardwoods mixed with bamboo, shrubs, and vines (Fig. 12). The air temperature was 29.2���36.5 o C and the relative humidity was 55���73%. Other species co-occurring with Opisthotropis voquyi include Cyclophiops multicinctus, Opisthotropis lateralis, Sinonatrix percarinata, Gekko palmatus, Sphenomorphus cryptotis, and Shinisaurus crocodilurus vietnamensis., Published as part of Ziegler, Thomas, David, Patrick, Ziegler, Tim N., Pham, Cuong T., Nguyen, Truong Q. & Le, Minh D., 2018, Morphological and molecular review of Jacob's Mountain Stream Keelback Opisthotropis jacobi Angel & Bourret, 1933 (Squamata: Natricidae) with description of a sibling species from northern Vietnam, pp. 476-496 in Zootaxa 4374 (4) on pages 481-494, DOI: 10.11646/zootaxa.4374.4.2, http://zenodo.org/record/1156694, {"references":["Angel, F. & Bourret, R. (1933) Sur une petite collection de serpents du Tonkin. Descriptions d'especes nouvelles. Bulletin de la Societe Zoologique de France, 58 (3 - 4), 129 - 140.","Stuart, B. L. & Chuaynkern, Y. (2007) A new Opisthotropis (Serpentes: Colubridae: Natricinae) from Northeastern Thailand. Current Herpetology, 26 (1), 35 - 40.","David, P., Pham, C. T., Nguyen, T. Q. & Ziegler, T. (2011) A new species of the genus Opisthotropis Gunther, 1872 (Squamata: Natricidae) from the highlands of Kon Tum Province, Vietnam. Zootaxa, 2758, 43 - 56.","Teynie, A., Lottier, A., David, P., Nguyen, T. Q. & Vogel, G. (2013) A new species of the genus Opisthotropis Gunther, 1872 from northern Laos (Squamata: Natricidae). Zootaxa, 3774 (2), 165 - 182. http: // dx. doi. org / 10.11646 / zootaxa. 3774.2.4","Wang, Y. Y., Guo, Q., Liu, Z. Y., Lyu, Z. T., Wang, J., Luo, L., Sun, Y. J. & Zhang, Y. W. (2017) Revisions of two poorly known species of Opisthotropis Gunther, 1872 (Squamata: Colubridae: Natricinae) with description of a new species from China. Zootaxa, 4247 (4), 391 - 412. https: // doi. org / 10.11646 / zootaxa. 4247.4.3","Hecht, V. L., Pham, C. T., Nguyen, T. T., Nguyen, T. Q., Bonkowski, M. & Ziegler, T. (2013) First Report on the herpetofauna of Tay Yen Tu Nature Reserve, northeastern Vietnam. Biodiversity Journal, 4 (4), 507 - 552.","Yang, J. H., Sung, Y. H. & Chan, B. P. L. (2013) A new species of the genus Opisthotropis Gunther, 1872 (Squamata: Colubridae: Natricinae) from Guangdong Province, China. Zootaxa, 3646 (3), 289 - 296."]}

Ziegler, Thomas, David, Patrick, Ziegler, Tim N., Pham, Cuong T., Nguyen, Truong Q., Le, Minh D. (2018): Morphological and molecular review of Jacob's Mountain Stream Keelback Opisthotropis jacobi Angel & Bourret, 1933 (Squamata: Natricidae) with description of a sibling species from northern Vietnam. Zootaxa 4374 (4): 476-496, DOI: https://doi.org/10.11646/zootaxa.4374.4.2

Opisthotropis voquyi sp. nov. Holotype. IEBR 4326 [Field No. TYT.2013.4], an adult male, from Tay Yen Tu Nature Reserve, Bac Giang Province, 437 m asl., Vietnam, collected by Cuong The Pham et al. on 14 June 2013 (Figs. 6–8). Paratypes. VNMN 0 6315 [Field No. TYT 2013.1], male, collected by Cuong The Pham et al. in June 2013; VNMN 0 6316 [Field No. BG 2015.13], female, collected by Cuong The Pham et al. on 17 May 2015; IEBR 4327 [Field No. BG 2015.33], female, collected by Cuong The Pham et al. on 22 May 2015; IEBR 4328 [Field No. BG 2015.34], female, collected by Cuong The Pham et al. on 22 May 2015; ZFMK 100819 [Field No. BG 2015.35], female, collected by Cuong The Pham et al. on 22 May 2015; ZFMK 100820 [Field No. TYT 2014.11], female, collected by Hang An Thi in May 2014, all paratypes same locality data as the holotype (Figs. 8–10). Bourret & Pa) Sa Bourret & Pa) Sa Chapa () 1934 a? Tamdao) b Chapa) (a 1935 Chapa (a 1935) Chapa a (1935) Tam Dao (1935) b Dao Tam (b) 1935 NganSon c1935 ()) (1936 Angel 1933 (syntype Angel (1933 syntype. 259 M Bourret A. 57 1934 (. M 518 Bourret 953 X. Bourret. 954 X Bourret 451 M. Bourret 794. X Bourret M. 498 Bourret Bourret ...Continued on next page TABLE 2b. (Continued) Diagnosis. A species of the genus Opisthotropis, characterized by a combination of the following characters: (1) internasal not in contact with loreal; (2) one preocular; (3) usually two postoculars; (4) one anterior temporal; (5) one posterior temporal; (6) 7 or 8, rarely 9 supralabials; (7) 25 maxillary teeth; (8) subcaudals 74–86; (9) 15 dorsal scale rows at neck, at midbody and before vent; (10) body scales smooth or only with few faint keels; (11) dorsal scales being grey, greyish-brown or brown in preservative, posteriorly more or less edged with pale greyishbrown. Description of holotype. Body stout, oval in cross section; head short, slightly broader than neck, depressed, dorsally covered with large shields; nostrils in dorsal position and oblique, narrow, piercing the middle of nasal, which is divided beneath; eye small, pupil round. Size. SVL: 337 mm; TaL: 106 mm; TL: 443 mm; ratio TaL/TL: 0.239. Dentition. Maxillary teeth: right upper maxilla with 25 subequal teeth or sockets, without diastema. Body scalation. Dorsal scale rows 15–15–15, smooth, with few faint keels. 181 ventrals (+ 1 preventral); 85 subcaudals, all paired; precloacal divided. Head scalation. Rostral heptagonal, wider than high, readily visible from above; nasals large, much longer than high, divided below nostril by a distinct furrow; internasals two, strongly curved, longer than wide, in contact with rostral, nasals, and prefrontal; prefrontal single, anteriorly pointed, much broader than long, in contact with internasals, nasals, loreals, preoculars, and frontal; frontal pentagonal, slightly wider than long, apex directed posteriorly, 1.3 times longer than prefrontal; parietals longer than wide, in contact approximately the length of frontal; 1 / 1 supraocular, distinctly wider than high; 1 / 1 loreal, pentagonal, slightly wider than high, not in contact with internasal; 1 / 1 preocular, large, hexagonal, distinctly higher than wide, reaching frontal, in broad contact with prefrontal; 2 / 2 postoculars, upper largest, curved, higher than wide, lower distinctly wider than high; 8 / 7 supralabials, anterior ones distinctly higher than long, fourth entering orbit (the fourth supralabial on the right side fused with the subsequent supralabial), SL 7 / 6 largest; 1+1 / 1+1 temporals, anterior one very long and narrow, in broad contact with SL 6–7 / 5–6 and parietals, posterior one more strongly developed; infralabials 9 / 9, first pair in contact behind small mental, IL 1–4 in contact with anterior chin shields, IL 5 largest, apex directed posteriorly; posterior chin shields about the same length than anterior ones, separated from each other by 2 + 4 scales. Hemipenes. Only the left hemipenis fully everted, ending in two short, inwards curved lobes (Fig. 7). Sulcus spermaticus undivided, extending only to the slightly smaller lobe. Hemipenes covered with small spines, curved backwards. Only the terminal area of the hemipenes between the lobes without spines. Upper truncus of hemipenes with ring of enlarged spines. Largest spines extending from sulcus spermaticus to hemipenis sides, with the lowermost one being the most distinct. Lower truncus of hemipenes covered with tiny spines. Coloration (in preservative). Upper head and body surface brown, the venter yellowish-beige. Dorsal scales posteriorly more or less edged with pale brown. Dorsal head surface paler around nostrils and in the middle of parietals. Infralabials same colour as dorsal scales, adjoining scales becoming gradually paler towards mental groove. Outermost edges of ventrals and subcaudals the same colour as dorsal scales. Subcaudals speckled with small dark spots, becoming more numerous towards the tail tip. Variation of paratypes. An overview of the meristic and scalation data of the type series of Opisthotropis voqyui sp. nov. is given in Table 3. The dorsal ground coloration varies between greyish, greyish brown, and brown. Concerning a potential sexual dimorphism, males seem to be brown, females grey or greyish brown. Females also seem to grow larger. The dorsal tail scales in the males had few faint keels, which was only the case for one female; the remaining four females had smooth dorsal tail scales. The paratype series generally accorded well with the holotype description. Of the type series six individuals had two postoculars, only one individual had 2 postoculars on the left side and one postocular on the right side. Comparisons. The comparison of Opisthotropis voquyi sp. nov. with other species of the genus Opisthotropis and the genera Parahelicops and Paratapinophis was based on the following references: Stuart & Chuaynkern (2007), David et al. (2011), Teynié et al. (2013), Nguyen et al. (in press), and Wang et al. (2017). There are only few species of Opisthotropis and other genera with 15 midbody dorsal scale rows (see Table 4). 15 15 or 17 17 19 23 Opisthotropis alcalai Brown & Leviton, 1961 x O. andersonii (Boulenger, 1888) x O. atra Günther, 1872 x O. balteata (Cope, 1895) x O. cheni Zhao, 1999 x O. cucae David, Pham, Nguyen & Ziegler, 2011 x O. daovantieni Orlov, Darevsky & Murphy, 1998 x O. durandi Teynié, Lottier, David, Nguyen & Vogel, 2013 x O. guangxiensis Zhao, Jiang & Huang, 1978 x O. jacobi Angel & Bourret, 1933 x O. kikuzatoi Okada & Takara, 1958 x O. kuatunensis Pope, 1928 x O. lateralis Boulenger, 1903 x O. latouchii (Boulenger, 1899) x O. laui Yang, Sung & Chan, 2013 x ...Continued on next page Main additional differences between Opisthotropis voquyi sp. nov. and Parahelicops annamensis are: one preocular in the new species versus two in P. annamensis; internasal not in contact with loreal in the new species versus in contact in P. annamensis; body scales smooth or only with few faint keels in the new species versus keeled throughout in P. annamensis and especially strongly keeled on the posterior part of the body and the base of the tail; body dorsum grey to greyish-brown or brown, with dorsals posteriorly more or less edged with pale greyish-brown in the new species versus dorsum with orange bars and spots in P. annamensis. Opisthotropis voquyi sp. nov. differs from O. guangxiensis by 7–9 versus 9–10 supralabials, one posterior temporal (versus two), 74–86 versus 51–58 subcaudals, dorsal scales arranged in 15–15–15 versus 17–15–15 rows, and a greyish to greyish–brown or brown dorsum, dorsal scales posteriorly more or less edged with pale greyishbrown in the new species versus dark body with pale crossbars in O. guangxiensis. Opisthotropis voquyi sp. nov. differs from O. kikuzatoi by 7–9 supralabials (versus 6), one preocular (versus two), one anterior temporal (versus two), and the greyish to greyish-brown or brown dorsal pattern, with the dorsals posteriorly being more or less edged with pale greyish-brown in the new species (versus dark dorsum with dorsolateral orange spots). Opisthotropis voquyi sp. nov. differs from O. maculosa by a grey to greyish-brown or brown dorsum, with dorsals being posteriorly more or less edged with pale greyish-brown in the new species versus with yellow dots on a dark background in O. maculosa. In external morphological characters, O. voquyi sp. nov. is particularly similar to O. jacobi, from which it differs in usually having two postoculars versus only one in O. jacobi, in the number of maxillary teeth: 25 in the new species versus 19–23 in O. jacobi, and in the dorsal scales being grey to greyish–brown or brown versus blackish–brown in O. jacobi. Etymology. The specific epithet is a noun in the genitive case. This species is named in honour of Professor Doctor Vo Quy of Vietnam National University, Hanoi, who passed away on 10 January 2017, and who played a pioneering role in nature conservation in Vietnam. He inspired later generations of conservation biologists, including authors of this study. Suggested common names. Vo Quy’s Mountain Stream Keelback (English) and Rắn trán võ quý (Vietnamese). Distribution. Opisthotropis voquyi is currently known only from the type locality in Bac Giang Province (Fig. 11). Natural history. Specimens were found at night between 19:30 and 23:00 h in and around small rocky streams, where the snakes were observed in the water or on the rocks close to the water. The surrounding habitat was secondary evergreen forest of medium and small hardwoods mixed with bamboo, shrubs, and vines (Fig. 12). The air temperature was 29.2–36.5 o C and the relative humidity was 55–73%. Other species co-occurring with Opisthotropis voquyi include Cyclophiops multicinctus, Opisthotropis lateralis, Sinonatrix percarinata, Gekko palmatus, Sphenomorphus cryptotis, and Shinisaurus crocodilurus vietnamensis.

Thomas Ziegler, Tim N. Ziegler, Julia Peusquens, Patrick David, Thanh N. Vu, Cuong T. Pham, Truong Q. Nguyen, Minh D. Le (2017): Expanded morphological definition and molecular phylogenetic position of the Tam Dao mountain stream keelback Opisthotropis tamdaoensis (Squamata: Natricidae) from Vietnam. Revue suisse de Zoologie 124 (2): 377-389, DOI: 10.5281/zenodo.893551, {"references":["Boulenger G.A. 1903. Descriptions of new snakes in the collection of the British Museum. Annals and Magazine of Natural History (7) 12: 350-354","Bourret R. 1936. Les serpents de l'Indochine. II. Catalogue systematique descriptif. Henri Basuyau et Cie, Toulouse, 505 pp.","Bui M.Q., Nguyen A.M.T., von Haeseler A. 2013. Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution 30: 1188-1195.","Burbrink F.T., Lawson R., Slowinski J.B. 2000. Mitochondrial DNA phylogeography of the polytypic North American rat snake (Elaphe obsoleta): a critique of the subspecies concept. Evolution 54: 2107-2118.","David P., Pham C.T., Nguyen T.Q., Ziegler T. 2011. A new species of the genus Opisthotropis Gunther, 1872 (Squamata: Natricidae) from the highlands of Kon Tum Province, Vietnam. Zootaxa 2758: 43-56.","Dowling H.G. 1951. A proposed standard system of counting ventrals in snakes. British Journal of Herpetology 1: 97-99.","Fan T.-H. 1931. Preliminary report of reptiles from Yaoshan, Kwangsi, China. Bulletin Department of Biology Sun Yatsen University 11: 1-154.","Gawor A., Pham C.T., Nguyen T.Q., Nguyen T.T., Schmitz A., Ziegler T. 2016. The herpetofauna of the Bai Tu Long National Park, northeast Vietnam. Salamandra 52(1): 23-41.","Guo P., Liu Q., Xu Y., Jiang K., Hou M., Ding L., Pyron R.A., Burbrink F.T. 2012. Out of Asia: natricine snakes support the Cenozoic Beringian Dispersal Hypothesis. Molecular Phylogenetics and Evolution 63(3): 825-833.","Hecht V.L., Pham C.T., Nguyen T.T., Nguyen T.Q., Bonkowski M., Ziegler T. 2013. First Report on the herpetofauna of Tay Yen Tu Nature Reserve, northeastern Vietnam. Biodiversity Journal 4(4): 507-552.","Le M., Raxworthy C.J., McCord W.P., Mertz L. 2006. A molecular phylogeny of tortoises (Testudines: Testudinidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 40: 517-531.","Mell R. 1930. Beitrage zur Lurch- und Kriechtierfauna Kwangsi's. 4. Schlangen. Sitzungsberichte Gesellschaft Naturfreunde Berlin 1929: 319-326.","Nguyen S.V., Ho C.T., Nguyen T.Q. 2009. Herpetofauna of Vietnam. Edition Chimaira, Frankfurt am Main, 768 pp.","Nguyen T.V., Nguyen T.Q., Pham C.T., Ziegler T. 2017. First country record of Opisthotropis maculosa Stuart & Chuaynkern, 2007 from Vietnam. Russian Journal of Herpetology (in press).","Nguyen L.T., Schmidt H. A., von Haeseler A., Bui M.Q. 2015. IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution 32: 268-274.","Pope C.H. 1935. The reptiles of China. Turtles, crocodilians, snakes, lizards. Natural History of Central Asia. Vol. X. American Museum of Natural History, New York, 604 pp.","Posada D., Crandall K.A. 1998. MODELTEST: testing the model of DNA substitution. Bioinformatics 14: 817-818.","Ronquist F., Teslenko M., van der Mark P., Ayres D.L., Darling A., Hohna S., Larget B., Liu L., Suchard M.A., Huelsenbeck J.P. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539-542.","Smith M.A. 1943. The fauna of British India, including Ceylon and Burma. Reptilia and Amphibia. Vol. III. Serpentes. Taylor & Francis Ltd., London, xii, 583 pp.","Stuart B.L., Chuaynkern Y. 2007. A new Opisthotropis (Serpentes: Colubridae: Natricinae) from Northeastern Thailand. Current Herpetology 26(1): 35-40.","Swofford D.L. 2001. PAUP*. Phylogenetic Analysis Using Parsimony (* and Other Methods), version 4. Sinauer Associates, Sunderland, Massachusetts.","Teynie A., Lottier A., David P., Nguyen T.Q., Vogel G. 2013. A new species of the genus Opisthotropis Gunther, 1872 from northern Laos (Squamata: Natricidae). Zootaxa 3774(2): 165-182.","Thompson J.D., Gibson T.J., Plewniak F., Jeanmougin F., Higgins D.G. 1997. The ClustalX windows interface: Flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 25: 4876-4882.","van Schingen M., Le M.D., Ngo H.T., Pham C.T., Ha Q.Q., Nguyen T.Q., Ziegler T. 2016. Is there more than one Crocodile Lizard? An integrative taxonomic approach reveals Vietnamese and Chinese Shinisaurus crocodilurus represent separate conservation and taxonomic units. Der Zoologische Garten, N.F. 85: 240-260.","Yang J.-H., Sung Y.-H., Chan B.P.-L. 2013. A new species of the genus Opisthotropis Gunther, 1872 (Squamata: Colubridae: Natricinae) from Guangdong Province, China. Zootaxa 3646(3): 289-296.","Ziegler T., David P., Thanh V. N. 2008. A new natricine snake of the genus Opisthotropis from Tam Dao, Vinh Phuc Province, northern Vietnam (Squamata, Colubridae). Zoosystematics and Evolution 84(2): 197-203.","Ziegler T., Rauhaus A., Tran T.D., Pham C.T., van Schingen M., Dang H.P. Le M.D., Nguyen T.Q. 2015. Die Amphibien- und Reptilienfauna der Me-Linh-Biodiversitatsstation in Nordvietnam. Sauria 37(4): 11-44."]}

Suwannapoom, Chatmongkon, Yuan, Zhi-Yong, Chen, Jin-Min, Hou, Mian, Zhao, Hai-Peng, Wang, Li-Jun, Nguyen, Truong Son, Nguyen, Truong Q., Murphy, Robert W., Sullivan, Jaqueline, Mcleod, David S., Che, Jing (2016): Taxonomic revision of the Chinese Limnonectes (Anura, Dicroglossidae) with the description of a new species from China and Myanmar. Zootaxa 4093 (2): 181-200, DOI: http://dx.doi.org/10.11646/zootaxa.4093.2.2

Leptolalax pallidus sp. nov. Figs. 8A, 11. Holotype. UNS 0 0 510, adult male, calling on plant leaf 0.1 m from ground, 1 m from 2–3 m wide rocky stream in evergreen forest, Bidoup Nui Ba National Park, Lam Dong Province, Vietnam. (12.1520 º N, 108.695º E; 1644 m elevation, Fig. 1, 9E). Collected at night on 27 June 2011 by Dao T. A. Tran and Duong T. T. Le. Paratypes. UNS 0 0 512, AMS R 177657, ZFMK 96599, three adult males collected on 26 July 2011 in in evergreen forest in Bidoup Nui Ba National Park, Lam Dong Province, Vietnam (12.1519º N, 108.6940º E; 1681 m elevation). UNS 0 0 511, AMS R 177659, ZFMK 96598, three adult males collected on same date and location as holotype. AMS R 177658, collected on 27 June 2011 at same location as holotype. All specimens were collected by Dao T. A. Tran and Duong T. T. Le Etymology. The specific name “ pallidus ”, meaning pale, is used as an adjective in reference to the paler coloration and less distinct markings of this species, particularly in preservative, compared to other species in the group. Diagnosis. Assigned to the genus Leptolalax on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra–axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax pallidus sp. nov. is distinguished from its congeners by a combination of (1) supra–axillary and ventrolateral glands present; (2) dark brownish red ventral surface with faint white speckling; (3) small SVL for the genus (24.5.1– 27.7 mm in 8 adult males), (4) toes lacking webbing and lateral fringes; (5) tibia length 45–51% of SVL in males; (6) pectoral gland 3.4–6.7% of SVL in males, (7) tuberculate skin texture on dorsum, (8) iris copper in upper half, gold in lower half; (9) distinct black supratympanic line absent, (10) an advertisement call with 4–7 notes with the short introductory note encompassing a quarter of each call and only one note in non-introductory notes. Description of holotype. Head width slightly subequal to head length; snout truncate in dorsal view and in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than to eye; canthus rostralis indistinct, gently rounded; lores sloping; pupil vertical; eye diameter smaller than snout length; tympanum indistinct, round, diameter smaller than that of the eye; tympanic rim slightly elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings small, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I Colour of holotype in life. Dorsal surface brown with slightly darker interorbital bar with pale brown patches anteriorly and posteriorly; faint dark W-shaped marking between axilla (Fig. 11A). Darker brown line along supratympanic ridge, terminating above axilla, encompassing most of tympanum; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms paler brown; numerous, small dark brown spots on sides from groin to axilla. Dark brownish ventral surface with white speckling on ventral surfaces of throat, chest and belly, more sparse on legs and arms. Supra-axillary gland copper; femoral glands white; pectoral glands white, ventrolateral glands white. Iris copper in upper half, gold in lower half, black reticulations throughout. Colour of holotype in preservative. Dorsum relatively uniform medium brown with darker interorbital bar and slightly paler posterior to this; dorsal surface of brachium, fingers and toes paler brown, dark brown/black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet (Fig. 8A). Elbows and upper arms pale brown. Ventral surface medium brown with faint white speckles. Macroglands creamy white. Measurements (mm). Holotype: SVL 25.2, HDL 9.4, HDW 9.2, SNT 3.9, EYE 3.4, IOD 3.2, TMP 1.7, TEY 1.4, TIB 13.0, EN 2.5, IN 2.6, NS 1.2, ML 6.8, PL 12.6. Variation. Measurements of the type series are shown in Tables 5–6 and representative photographs of paratypes are shown in Fig. 11. All specimens have a degree of darker brown patterning (mostly marbling but also some spots) on the dorsum and dark banding on the tibiotarsus and antebrachium, hands and feet, but both are more pronounced in UNS00512, AMS R 177657 and UNS00511. ZFMK 96599 has faint dark brown spots evenly distributed around tubercles on dorsum. UNS00510–00512 have more distinctly pale elbows in preservative. Size and distribution of white speckles on the venter varies a little between specimens, more clumped on chest in AMS R 177659 and ZFMK 96598. Skin texture less tuberculate in preservative than in life. Advertisement call. Call descriptions are based on the calls of three individuals, recorded at 14.0–21.4 ºC ambient temperature. Calls were an average of 673 ms in duration and consisted of 4–7 notes (Table 7, Fig. 6A). Short introductory notes of low relative amplitude were present in all calls. Introductory notes were 22–26% of the call duration, contained up to 63 pulses. Non-introductory notes invariably contained a single pulse. Amplitude peaked towards the middle of the call. Notes were repeated an average of 6.8 notes/s. The average dominant frequency was 2.5 kHz. Harmonics were weakly present at ~7.3 kHz, and a fundamental frequency was not visible. To the human ear, the advertisement call of L. pallidus sp. nov. consists of a squelch followed by clicking. Ecology. All specimens of Leptolalax pallidus sp. nov. were found in evergreen forest in Bidoup Nui Ba National Park between 1644–1681 m elevation. Males were located on the forest floor, on leaf-litter or rocky crevices and clay holes in the banks of a dry stream. Fig 9E. Distribution. Leptolalax pallidus sp. nov. is only known from Gia Rich mountain in Bidoup Nui Ba National Park, only about 5 km from L. bidoupensis. The two species occur in continuous forest, with no obvious current biogeographic barrier between them. Comparisons. Leptolalax pallidus sp. nov. differs from all other Leptolalax species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from L. aereus by ventral coloration and iris coloration, from L. bourreti by ventral coloration, male body size, absence of black supratympanic line, skin texture and iris coloration, from L. botsfordi by male body size, absence of black supratympanic line, iris coloration and relative male body weight (0.42–0.50 versus 0.75–0.94 g /mm), from L. croceus by ventral coloration and iris coloration; from L. eos by ventral coloration, male body size, iris coloration and lateral fringes on toes; from L. firthi by ventral coloration, absence of black supratympanic line, iris coloration and lateral fringes on toes; from L. fuliginosus by ventral coloration, male body size, skin texture and iris coloration; from L. heteropus by ventral coloration, male body size, skin texture and absence of black supratympanic line; from L. kecil by ventral coloration, male body size, absence of black supratympanic line and iris coloration; from L. melanoleucus by ventral coloration, body size, absence of black supratympanic line, skin texture and iris coloration; from L. minimus by ventral coloration, skin texture and male body size; from L. nahangensis by ventral coloration, male body size, skin texture, absence of black supratympanic line and iris coloration; from L. nyx by ventral coloration, skin texture and absence of black supratympanic line, from L pelodytoides by ventral coloration and absence of black supratympanic line; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration, body size, skin texture and absence of black supratympanic line; from L. solus by ventral coloration, absence of black supratympanic line, skin texture and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. ventripunctatus by ventral coloration, absence of black supratympanic line, skin texture and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size, skin texture and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. pallidus sp. nov. differs by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly larger and non-overlapping male body size (Wilcoxon post-hoc Z=3.4, pL. applebyi) and tuberculate skin texture (versus mostly smooth in L. applebyi). L. pallidus sp. nov. differs from L. bidoupensis by having a significantly larger body size (Wilcoxon post-hoc Z=2.7, p=0.006), larger relative tibia length (Wilcoxon post-hoc Z=3.0, p=0.002), larger relative tympanum size (Wilcoxon post-hoc Z=2.0, p=0.041), dark brown iris (versus reddish upper and silver below), indistinct, brown supratympanic line (versus distinct black supratypmanic line in L. bidoupensis) and tuberculate skin texture (versus mostly smooth in L. bidoupensis). L. pallidus sp. nov. differs from L. melicus by having a significantly larger and non-overlapping male body size (Wilcoxon post-hoc Z=-3.3, pL. applebyi) and tuberculate skin texture (versus mostly smooth in L. melicus). L. pallidus sp. nov. differs from L. pyrrhops by having a significantly smaller and non-overlapping male body size (Wilcoxon post-hoc Z=3.2, p=0.002), smaller relative diameter of the eye (Wilcoxon post-hoc Z=3.2, p=0.002), indistinct, brown supratympanic line (versus distinct black supratypmanic line in L. pyrrhops) and tuberculate skin texture (versus only slightly shagreened in L. pyrrhops). L. pallidus sp. nov. differs from L. ardens sp. nov. by having a significantly larger body size (Wilcoxon post-hoc Z=-3.8, pL. ardens sp. nov) and tuberculate skin texture (versus mostly smooth in L. ardens sp. nov). L. pallidus sp. nov. differs from L. kalonensis sp. nov. by having a significantly smaller body size (Wilcoxon post-hoc Z=3.0, p=0.002), larger relative tympanum diameter (Wilcoxon post-hoc Z=-3.1, p=0.002), indistinct, brown supratympanic line (versus distinct black supratypmanic line in most L. kalonensis sp. nov) and tuberculate skin texture (versus mostly smooth in L. kalonensis sp. nov). L. pallidus sp. nov. differs from L. maculosus sp. nov. by having indistinct, brown supratympanic line (versus distinct black supratypmanic line in L. maculosus sp. nov) and tuberculate skin texture (versus mostly smooth in L. maculosus sp. nov). L. pallidus sp. nov. differs from L. tadungensis sp. nov. by having a significantly larger relative tibia length (Wilcoxon post-hoc Z=-3.1, p=0.002), larger relative distance between the eye and tympanum (Wilcoxon post-hoc Z=-2.4, p=0.019), indistinct, brown supratympanic line (versus distinct black supratympanic line in L. maculosus sp. nov) and tuberculate skin texture (versus mostly smooth in L. maculosus sp. nov). See Table 4 and Fig. 4. The male advertisement call of L. pallidus sp. nov. differs from all members of the L. applebyi group. The call of L. pallidus sp. nov. differs from L. applebyi in call duration, having a relatively long, pulsed introductory note, having one pulse in non-introductory notes, and by average note repetition rate and average dominant frequency. L. pallidus sp. nov. differs from L. bidoupensis in call duration, in having a distinct, pulsed introductory note, and by average note repetition rate and perhaps average dominant frequency (non-overlapping but similar values); from L. melicus in call duration, number of pulses in the introductory note, average note repetition rate and average dominant frequency; from L. pyrrhops in call duration, average number of pulses in introductory note, average note repetition rate and potentially by dominant frequency; from L. ardens sp. nov. differs by call duration, number of pulses in the introductory note, note repetition rate and dominant frequency; from L. kalonensis sp. nov. in average call duration, average number of pulses in the introductory note and average note repetition rate; from L. maculosus sp. nov. in average call duration number of notes per call, and having a distinct introductory note; from L. tadungensis sp. nov. in average call duration, average duration of introductory note, and note repetition rate. See Table 7 and Figs. 5–6. Leptolalax pallidus sp. nov. differs from all species within the L. applebyi group by>4.7% divergence at the 16S gene fragment examined. Interspecific variation in three L. pallidus sp. nov. collected from

Rowley, Jodi J. L., Tran, Dao T. A., Le, Duong T. T., Dau, Vinh Q., Peloso, Pedro L. V., Nguyen, Truong Q., Hoang, Huy D., Nguyen, Tao T., Ziegler, Thomas (2016): Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam. Zootaxa 4085 (1): 63-102, DOI: http://doi.org/10.11646/zootaxa.4085.1.3

Leptolalax maculosus sp. nov. Figs. 8C, 9C, 13. Holotype. UNS 00513, adult male, in dry stream bed in evergreen forest in Phuoc Binh National Park, Ninh Thuan Province, Vietnam (12.0152º N, 108.7271º E, 900 m elevation, Fig. 1). Collected on 24 September 2011 by Dao T. A. Tran and Duong T. T. Le. Paratypes. ZFMK 96600, adult male from same date and location as holotype. AMS R 177660, adult male, and UNS 00514, adult female, collected on 27 September 2011 from evergreen forest in Phuoc Binh National Park, Ninh Thuan Province, Vietnam (12.0413º N, 108.7385º E, 1166 m elevation). All specimens were collected by Dao T. A. Tran and Duong T. T. Le. Etymology. The specific name “ maculosus ”, meaning spotted, speckled, mottled or variegated, is used as an adjective in reference to the more mottled dorsal pattern of this species. Diagnosis. Assigned to the genus Leptolalax on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra–axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax maculosus sp. nov. is distinguished from its congeners by a combination of (1) supra-axillary and ventrolateral glands present; (2) dark brownish red ventral surface with white speckling; (3) small/medium SVL for the genus (24.2–26.6 mm in three adult males, 27.0 mm in one female); (4) toes lacking webbing and lateral fringes; (5) tibia length 48–50% of SVL in males; (6) pectoral gland 3.5–4.2% of SVL in males, (7) mostly smooth dorsum, (8) iris copper in upper half and gold in lower half (9) distinct black supratympanic line present, (10) an advertisement call with 7–38 notes, lacking a distinct introductory note. Description of holotype. Head width equal to head length; snout rounded in dorsal view and slightly truncate in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than to eye; canthus rostralis indistinct, gently rounded; lores straight; pupil vertical; eye diameter slightly larger than snout length; tympanum distinct, round, diameter smaller than that of the eye; tympanic rim slightly elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings slit–like, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I Colour of holotype in life. Dorsal surface dark brown with irregular, darker brown blotch down back; small, pale brown blotches scattered over dorsum; dark interorbital bar with pale copper edging anteriorly; two small, pale copper V-markings between axilla (Fig. 13A, B). Black line along supratympanic ridge, terminating above axilla, encompassing most of tympanum; irregular copper band under black supratympanic line; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms copper; numerous, small darker brown spots on sides from groin to axilla. Brownish pink ventral surface with white speckling on ventral surfaces of throat, chest, belly, legs and arms. Supra-axillary gland copper; femoral glands white; pectoral glands white, ventrolateral glands white. Iris copper in upper half, gold in lower half, black reticulations throughout. Colour of holotype in preservative. Dorsum medium brown with small, pale brown blotches on top of head, barely visible paler intraorbital blotch; dark brown/black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet (Fig. 8C). Black blotch on groin. Ventral surface medium brown with white speckling. Macroglands creamy white. Measurements (mm). Holotype: SVL 24.2, HDL 8.9, HDW 9.0, SNT 3.3, EYE 3.4, IOD 3.5, TMP 1.5, TEY 0.9, TIB 11.6, EN 2.3, IN 2.3, NS 1.1, ML 6.0, PL 10.1. Variation: Measurements of the type series are shown in Tables 5–6 and representative photographs of paratypes are shown in Figs. 9C and 15. AMS R 177660 and to a lesser extent ZFMK 96600 have more distinct patterning, with paler blotches on the dorsum Advertisement call. Call descriptions are based on the calls of two individuals, recorded at 23.3–24.1 ºC ambient temperature. Calls were an average of 907 ms in duration and consisted of 7–38 notes (Table 7, Fig. 6C). Calls were highly amplitude modulated, with a slow rise in amplitude, and amplitude peaking towards the middle of each note. Notes near-invariably contained a single pulse. Notes were repeated an average of 17 notes/s, but note repetition rate was higher at the start of each call. The average dominant frequency was 2.7 kHz, with peak frequency spread over ~2.7–3.5 kHz. A fundamental frequency was not visible. To the human ear, the advertisement call of L. maculosus sp. nov. is a squealch grading gently into a chirp. Ecology. All specimens of Leptolalax maculosus sp. nov. were found in evergreen forest between 900–1166 m elevation at Phuoc Binh National Park. Specimens were found within leaf-litter and in clay holes in a dry stream bed of 1.5–2.0 m width, connecting to a larger stream. Distribution. Leptolalax maculosus sp. nov. is only known from Phuoc Binh National Park, Ninh Thuan Province, Vietnam. The maximum distance between known localities is only 3.2 km. Comparisons. Leptolalax maculosus sp. nov. differs from all other Leptolalax species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from L. aereus by ventral coloration and presence of distinct black supratympanic line, from L. bourreti by ventral coloration and male body size, from L. botsfordi by male body size and relative body weight (0.41–0.47 versus 0.75–0.94 g /mm in L. botsfordi), from L. croceus by ventral coloration, presence of black supratympanic line and iris coloration; from L. eos by ventral coloration, male body size, presence of black supratympanic line and lateral fringes on toes; from L. firthi by ventral coloration, iris coloration and lateral fringes on toes; from L. fuliginosus by ventral coloration, male body size and iris coloration; from L. heteropus by ventral coloration and iris coloration; from L. kecil by ventral coloration, male body size and iris coloration; from L. melanoleucus by ventral coloration and iris coloration; from L. minimus by ventral coloration; from L. nahangensis by ventral coloration and male body size; from L. nyx by ventral coloration and male body size, from L pelodytoides by ventral coloration and male body size; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration and male body size; from L. solus by ventral coloration, male body size and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, presence of black supratympanic line and skin texture; from L. ventripunctatus by ventral coloration and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. maculosus sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly greater body size (Wilcoxon post-hoc Z=2.7, p=0.007), greater relative head width (Wilcoxon post-hoc Z=2.7, p=0.007), greater relative diameter of the eye (Wilcoxon post-hoc Z=2.2, p=0.025), and greater relative pectoral gland size (Wilcoxon post-hoc Z=2.4, p=0.017). L. maculosus sp. nov. differs from L. bidoupensis by having a significantly greater relative tibia length (Wilcoxon post-hoc Z=2.4, p=0.018), and a bronze iris (versus reddish upper and silver below in L. bidoupensis). L. maculosus sp. nov. differs from L. melicus by having a significantly greater body size (Wilcoxon post-hoc Z=2.6, p=0.009), and greater length between eye and tympanum (Wilcoxon post-hoc Z=2.1, p=0.034). L. maculosus sp. nov. differs from L. pyrrhops by having a significantly smaller body size (Wilcoxon post-hoc Z= -2.6, p=0.010), smaller relative length between eye and tympanum (Wilcoxon post-hoc Z= -2.4, p=0.018) and smaller relative diameter of the eye (Wilcoxon posthoc Z=-2.4, p=0.018). L. maculosus sp. nov. differs from L. ardens sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z=2.8, p=0.006). L. maculosus sp. nov. differs from L. pallidus sp. nov. by having a black supratympanic line (versus no black supratypmanic line in L. pallidus sp. nov) and smooth- finely shagreened skin texture (versus tuberculate in L. pallidus sp. nov). L. maculosus sp. nov. differs from L. kalonensis sp. nov. by having a significantly smaller body size (Wilcoxon post-hoc Z=-2.7, p=0.007). L. maculosus sp. nov. differs from L. tadungensis sp. nov. by having a significantly greater relative tibia length (Wilcoxon post-hoc Z=-2.5, p=0.013). See Table 4 and Fig. 5. The male advertisement call of L. maculosus sp. nov. differs from all species in the L. applebyi group. The call of the new species differs from L. applebyi in call duration, number of notes per call, an introductory note, and dominant frequency; from L. bidoupensis in call duration, average number of notes per call, presence of an introductory note, and average dominant frequency; from L. melicus in call duration, number of notes per call, having an indistinct introductory note, and dominant frequency; from L. pyrrhops by call duration, number of notes per call, having an indistinct introductory note, and dominant frequency; from L. ardens sp. nov. by call duration, number of notes per call and dominant frequency; from L. pallidus sp. nov. by call duration and number of notes per call; from L. kalonensis sp. nov. by average call duration, and number of notes per call; from L. tadungensis sp. nov. by call duration, number of notes per call, having a less distinct introductory note and different relative duration of introductory note. See Table 7 and Figs. 5–6. Leptolalax maculosus sp. nov. differs from all species within the L. applebyi group by>6.3% divergence at the 16S gene fragment examined. Interspecific variation in four L. maculosus sp. nov. collected from up to ~ 3 km apart was 0.0–0.2%.

Leptolalax tadungensis sp. nov. Figs. 8D, 14. Holotype. UNS 00516, adult male, in dry stream in evergreen forest in Ta Dung Nature Reserve, Dak Nong Province, Vietnam (11.8350º N, 108.0081º E, 1262 m elevation, Fig. 1, 9G). Collected on 13 July 2010 by Dao T. A. Tran, Vinh Q. Dau, and Huy D. Hoang. Paratypes. AMS R 177666 and ZFMK 96601, two adult males, collected on same date and locality as holotype. UNS00515, AMS R 177661, AMS R 177662, ZFMK 96602 and ZFMK 96603, five adult males collected on 17 July 2010 in Ta Dung Nature Reserve, Dak Nong Province, Vietnam (11.8388º N, 107.8945º E, 720 m elevation). UNS 0 0 526 and UNS 0 0 527, two adult males, collected on 17 July 2011 in Ta Dung Nature Reserve, Dak Nong Province, Vietnam (11.8561º N, 108.0371º E, 1932 m elevation); UNS 0 0 517 adult female, collected at same site on 16 July 2011. Specimens were collected by Dao T. A. Tran and Huy D. Hoang. Etymology. Specific epithet “ tadungensis ” is in reference to the type locality of the species, Ta Dung Nature Reserve. Diagnosis. Assigned to the genus Leptolalax on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra–axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax tadungensis sp. nov. is distinguished from its congeners by a combination of (1) supra–axillary and ventrolateral glands present; (2) dark brownish red ventral surface with white speckling; (3) small/medium SVL for the genus (23.5–26.0 mm in eight adult males, 32.1 mm in one female); (4) toes lacking webbing and lateral fringes; (5) tibia length 43–48% of SVL in males; (6) pectoral gland 3.3–5.4% of SVL in males, (7) smooth dorsum, (8) iris dark gold, (9) distinct black supratympanic line present, (10) an advertisement call with 4–7 notes with the first, introductory note encompassing less than a tenth of each call. Description of holotype. Head width less than head length; snout rounded in dorsal view and slightly truncate in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than eye; canthus rostralis indistinct, gently rounded; lores straight; vertical pupil; eye diameter smaller than snout length; tympanum distinct, round, diameter smaller than that of the eye; tympanic rim slightly elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings slit–like, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I Colour of holotype in life. Dorsal surface dark brown; very faint transverse darker brown bars on the dorsal surface of the thighs, tibia, tarsus, lower arms, fingers and toes. Black line along supratympanic ridge, terminating above axilla, encompassing posterodorsal half of tympanum; anteroventral half paler brown; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms slightly paler, brownish copper; numerous, small darker brown spots on sides from groin to axilla. Dark brown ventral surface with fine white speckling on ventral surfaces of throat, chest, belly, legs and arms; most dense on chest. Supraaxillary gland copper; femoral glands white; pectoral glands white. Iris copper, fine black reticulations throughout. Colour of holotype in preservative. Dorsum uniform dark brown with scattered small black blotches on flanks and larger black spot on groin; fine black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet (Fig. 8D). Ventral surface medium brown with only very faintly visible, fine white speckling. Macroglands white. Measurements (mm). Holotype: SVL 24.9, HDL 9.4, HDW 8.9, SNT 3.8, EYE 2.9, IOD 3.4, TMP 1.6, TEY 0.9, TIB 11.7, EN 2.2, IN 2.2, NS 1.7, ML 6.0, PL 10.4. Variation. Measurements of the type series are shown in Tables 5–6 and representative photographs of paratypes are shown in Fig. 14. Little variation in dorsal colouration. Single female specimen UNS 0 0 527 is paler in colour with a more truncate snout. The white speckling on venter is more distinct in UNS 0 0 527, UNS 00517, and AMS R 177661, forming small blotches in UNS 0 0 527, and UNS 00517; the dorsolateral line is more distinct and less broken in these specimens. UNS 0 0 526 and UNS 00517 have a slightly shagreened skin texture in preservative, possibly an artefact of preservation (with these specimens being better preserved). Advertisement call. Call descriptions are based on the calls of four individuals, recorded at 12.9–22.3ºC ambient temperature. Calls were an average of 291 ms in duration and consisted of 4–7 notes (Table 7, Figure 6D). Introductory notes were very short and quite similar to non-introductory notes, comprising only 3–9% of the call duration, and were indistinctly pulsed. Non-introductory notes contained 1–4 pulses. Notes were repeated an average of 14.3 notes/s. The average dominant frequency was 2.8 kHz. Harmonics were weakly present at ~6.3, 13.2, 16.6 kHz, and a fundamental frequency was not visible. To the human ear, the advertisement call of L. pallidus sp. nov. is a short raspy chirp, similar to an orthopteran. Ecology. All specimens of Leptolalax tadungensis sp. nov. were found in evergreen forest in Ta Dung Nature Reserve between 720–1932 m elevation (Fig. 9H). All frogs were found calling during rainy conditions. Frogs collected at the type locality were in underground rocky crevices. Individuals collected on Ta Dung Mountain (~ 1930 m elevation) were calling from the ground or under leaf litter. Distribution. Leptolalax tadungensis sp. nov. is only known from Ta Dung Nature Reserve, Dak Nong Province, Vietnam. The greatest distance between known localities for the species is 15.7 km. Comparisons. Leptolalax tadungensis sp. nov. differs from all other Leptolalax species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from L. aereus by ventral coloration and presence of distinct black supratympanic line, from L. bourreti by ventral coloration, from L. botsfordi by male body size, from L. croceus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. eos by ventral coloration, male body size, presence of black supratympanic line and lateral fringes on toes; from L. firthi by ventral coloration, iris coloration, skin texture and lateral fringes on toes; from L. fuliginosus by ventral coloration and iris coloration; from L. heteropus by ventral coloration and iris coloration; from L. kecil by ventral coloration, male body size and iris coloration; from L. melanoleucus by ventral coloration and iris coloration; from L. minimus by ventral coloration; from L. nahangensis by ventral coloration and male body size; from L. nyx by ventral coloration; from L pelodytoides by ventral coloration; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration and male body size; from L. solus by ventral coloration and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, presence of black supratympanic line and skin texture; from L. ventripunctatus by ventral coloration and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. tadungensis sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly greater body size (Wilcoxon post-hoc Z=3.6, p=0.001), greater relative diameter of the eye (Wilcoxon post-hoc Z=2.5, p=0.013), and greater relative pectoral gland size (Wilcoxon posthoc Z=2.9, p=0.004). L. tadungensis sp. nov. differs from L. bidoupensis by having a significantly smaller relative head width (Wilcoxon post-hoc Z=-2.1, p=0.032), smaller relative distance between eye and tympanum (Wilcoxon post-hoc Z=-2.1, p=0.032), and gold iris (versus reddish upper and silver below in L. bidoupensis). L. tadungensis sp. nov. differs from L. melicus by having a significantly larger body size (Wilcoxon post-hoc Z=3.5, p=0.001), and smaller relative tibia length (Wilcoxon post-hoc Z= -2.3, p=0.019). L. tadungensis sp. nov. differs from L. pyrrhops by having a significantly smaller body size (Wilcoxon post-hoc Z= -3.4, p=0.001), smaller relative tibia length (Wilcoxon post-hoc Z= -3.2, p=0.002), smaller relative distance between eye and tympanum (Wilcoxon post-hoc Z=-3.1, p=0.002) and smaller relative diameter of the eye (Wilcoxon post-hoc Z= -3.3, p=0.001). L. tadungensis sp. nov. differs from L. ardens sp. nov. by having a significantly larger body size (Wilcoxon post-hoc Z= 3.4, p=0.001) and smaller relative tibia length (Wilcoxon post-hoc Z= -3.0, p=0.003). L. tadungensis sp. nov. differs from L. pallidus sp. nov. by having smaller relative tibia length (Wilcoxon post-hoc Z= -3.1, p=0.002), smaller relative distance between eye and tympanum (Wilcoxon post-hoc Z= -2.4, p=0.019), a black supratympanic line (versus no black supratypmanic line in L. pallidus sp. nov) and mostly smooth skin texture (versus tuberculate in L. pallidus sp. nov). L. tadungensis sp. nov. differs from L. kalonensis sp. nov. by having a significantly smaller body size (Wilcoxon post-hoc Z= -3.3, p=0.001) and smaller relative tibia length (Wilcoxon post-hoc Z= -3.4, p=0.001). L. tadungensis sp. nov. differs from L. maculosus sp. nov. by having a significantly smaller relative tibia length (Wilcoxon post-hoc Z= -2.5, p=0.013). See Table 4 and Fig. 4. The male advertisement call of L. tadungensis sp. nov. differs from all species in the L. applebyi group. The call of the new species differs from L. applebyi in dominant frequency; from L. bidoupensis in having an introductory note present and in dominant frequency; from L. melicus in relative length of introductory note and dominant frequency; from L. pyrrhops by dominant frequency; from L. ardens sp. nov. by relative length of introductory note; from from L. pallidus sp. nov. by call duration and relative length of introductory note; from L. kalonensis sp. nov. by relative length of introductory note; from L. maculosus sp. nov. by call duration, number of notes per call, distinctness of introductory note and different relative duration of introductory note. See Table 7 and Fig. 5–6. Leptolalax tadungensis sp. nov. differs from all species within the L. applebyi group by>6.3% divergence at the 16S gene fragment examined. Interspecific variation in four L. tadungensis sp. nov. collected from up to ~ 15 km apart was 0.0–0.2%.

Leptolalax maculosus sp. nov. Figs. 8C, 9C, 13. Holotype. UNS 00513, adult male, in dry stream bed in evergreen forest in Phuoc Binh National Park, Ninh Thuan Province, Vietnam (12.0152�� N, 108.7271�� E, 900 m elevation, Fig. 1). Collected on 24 September 2011 by Dao T. A. Tran and Duong T. T. Le. Paratypes. ZFMK 96600, adult male from same date and location as holotype. AMS R 177660, adult male, and UNS 00514, adult female, collected on 27 September 2011 from evergreen forest in Phuoc Binh National Park, Ninh Thuan Province, Vietnam (12.0413�� N, 108.7385�� E, 1166 m elevation). All specimens were collected by Dao T. A. Tran and Duong T. T. Le. Etymology. The specific name ��� maculosus ���, meaning spotted, speckled, mottled or variegated, is used as an adjective in reference to the more mottled dorsal pattern of this species. Diagnosis. Assigned to the genus Leptolalax on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra���axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax maculosus sp. nov. is distinguished from its congeners by a combination of (1) supra-axillary and ventrolateral glands present; (2) dark brownish red ventral surface with white speckling; (3) small/medium SVL for the genus (24.2���26.6 mm in three adult males, 27.0 mm in one female); (4) toes lacking webbing and lateral fringes; (5) tibia length 48���50% of SVL in males; (6) pectoral gland 3.5���4.2% of SVL in males, (7) mostly smooth dorsum, (8) iris copper in upper half and gold in lower half (9) distinct black supratympanic line present, (10) an advertisement call with 7���38 notes, lacking a distinct introductory note. Description of holotype. Head width equal to head length; snout rounded in dorsal view and slightly truncate in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than to eye; canthus rostralis indistinct, gently rounded; lores straight; pupil vertical; eye diameter slightly larger than snout length; tympanum distinct, round, diameter smaller than that of the eye; tympanic rim slightly elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings slit���like, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I Colour of holotype in life. Dorsal surface dark brown with irregular, darker brown blotch down back; small, pale brown blotches scattered over dorsum; dark interorbital bar with pale copper edging anteriorly; two small, pale copper V-markings between axilla (Fig. 13A, B). Black line along supratympanic ridge, terminating above axilla, encompassing most of tympanum; irregular copper band under black supratympanic line; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms copper; numerous, small darker brown spots on sides from groin to axilla. Brownish pink ventral surface with white speckling on ventral surfaces of throat, chest, belly, legs and arms. Supra-axillary gland copper; femoral glands white; pectoral glands white, ventrolateral glands white. Iris copper in upper half, gold in lower half, black reticulations throughout. Colour of holotype in preservative. Dorsum medium brown with small, pale brown blotches on top of head, barely visible paler intraorbital blotch; dark brown/black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet (Fig. 8C). Black blotch on groin. Ventral surface medium brown with white speckling. Macroglands creamy white. Measurements (mm). Holotype: SVL 24.2, HDL 8.9, HDW 9.0, SNT 3.3, EYE 3.4, IOD 3.5, TMP 1.5, TEY 0.9, TIB 11.6, EN 2.3, IN 2.3, NS 1.1, ML 6.0, PL 10.1. Variation: Measurements of the type series are shown in Tables 5���6 and representative photographs of paratypes are shown in Figs. 9C and 15. AMS R 177660 and to a lesser extent ZFMK 96600 have more distinct patterning, with paler blotches on the dorsum Advertisement call. Call descriptions are based on the calls of two individuals, recorded at 23.3���24.1 ��C ambient temperature. Calls were an average of 907 ms in duration and consisted of 7���38 notes (Table 7, Fig. 6C). Calls were highly amplitude modulated, with a slow rise in amplitude, and amplitude peaking towards the middle of each note. Notes near-invariably contained a single pulse. Notes were repeated an average of 17 notes/s, but note repetition rate was higher at the start of each call. The average dominant frequency was 2.7 kHz, with peak frequency spread over ~2.7���3.5 kHz. A fundamental frequency was not visible. To the human ear, the advertisement call of L. maculosus sp. nov. is a squealch grading gently into a chirp. Ecology. All specimens of Leptolalax maculosus sp. nov. were found in evergreen forest between 900���1166 m elevation at Phuoc Binh National Park. Specimens were found within leaf-litter and in clay holes in a dry stream bed of 1.5���2.0 m width, connecting to a larger stream. Distribution. Leptolalax maculosus sp. nov. is only known from Phuoc Binh National Park, Ninh Thuan Province, Vietnam. The maximum distance between known localities is only 3.2 km. Comparisons. Leptolalax maculosus sp. nov. differs from all other Leptolalax species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from L. aereus by ventral coloration and presence of distinct black supratympanic line, from L. bourreti by ventral coloration and male body size, from L. botsfordi by male body size and relative body weight (0.41���0.47 versus 0.75���0.94 g /mm in L. botsfordi), from L. croceus by ventral coloration, presence of black supratympanic line and iris coloration; from L. eos by ventral coloration, male body size, presence of black supratympanic line and lateral fringes on toes; from L. firthi by ventral coloration, iris coloration and lateral fringes on toes; from L. fuliginosus by ventral coloration, male body size and iris coloration; from L. heteropus by ventral coloration and iris coloration; from L. kecil by ventral coloration, male body size and iris coloration; from L. melanoleucus by ventral coloration and iris coloration; from L. minimus by ventral coloration; from L. nahangensis by ventral coloration and male body size; from L. nyx by ventral coloration and male body size, from L pelodytoides by ventral coloration and male body size; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration and male body size; from L. solus by ventral coloration, male body size and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, presence of black supratympanic line and skin texture; from L. ventripunctatus by ventral coloration and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. maculosus sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly greater body size (Wilcoxon post-hoc Z=2.7, p=0.007), greater relative head width (Wilcoxon post-hoc Z=2.7, p=0.007), greater relative diameter of the eye (Wilcoxon post-hoc Z=2.2, p=0.025), and greater relative pectoral gland size (Wilcoxon post-hoc Z=2.4, p=0.017). L. maculosus sp. nov. differs from L. bidoupensis by having a significantly greater relative tibia length (Wilcoxon post-hoc Z=2.4, p=0.018), and a bronze iris (versus reddish upper and silver below in L. bidoupensis). L. maculosus sp. nov. differs from L. melicus by having a significantly greater body size (Wilcoxon post-hoc Z=2.6, p=0.009), and greater length between eye and tympanum (Wilcoxon post-hoc Z=2.1, p=0.034). L. maculosus sp. nov. differs from L. pyrrhops by having a significantly smaller body size (Wilcoxon post-hoc Z= -2.6, p=0.010), smaller relative length between eye and tympanum (Wilcoxon post-hoc Z= -2.4, p=0.018) and smaller relative diameter of the eye (Wilcoxon posthoc Z=-2.4, p=0.018). L. maculosus sp. nov. differs from L. ardens sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z=2.8, p=0.006). L. maculosus sp. nov. differs from L. pallidus sp. nov. by having a black supratympanic line (versus no black supratypmanic line in L. pallidus sp. nov) and smooth- finely shagreened skin texture (versus tuberculate in L. pallidus sp. nov). L. maculosus sp. nov. differs from L. kalonensis sp. nov. by having a significantly smaller body size (Wilcoxon post-hoc Z=-2.7, p=0.007). L. maculosus sp. nov. differs from L. tadungensis sp. nov. by having a significantly greater relative tibia length (Wilcoxon post-hoc Z=-2.5, p=0.013). See Table 4 and Fig. 5. The male advertisement call of L. maculosus sp. nov. differs from all species in the L. applebyi group. The call of the new species differs from L. applebyi in call duration, number of notes per call, an introductory note, and dominant frequency; from L. bidoupensis in call duration, average number of notes per call, presence of an introductory note, and average dominant frequency; from L. melicus in call duration, number of notes per call, having an indistinct introductory note, and dominant frequency; from L. pyrrhops by call duration, number of notes per call, having an indistinct introductory note, and dominant frequency; from L. ardens sp. nov. by call duration, number of notes per call and dominant frequency; from L. pallidus sp. nov. by call duration and number of notes per call; from L. kalonensis sp. nov. by average call duration, and number of notes per call; from L. tadungensis sp. nov. by call duration, number of notes per call, having a less distinct introductory note and different relative duration of introductory note. See Table 7 and Figs. 5���6. Leptolalax maculosus sp. nov. differs from all species within the L. applebyi group by>6.3% divergence at the 16S gene fragment examined. Interspecific variation in four L. maculosus sp. nov. collected from up to ~ 3 km apart was 0.0���0.2%., Published as part of Rowley, Jodi J. L., Tran, Dao T. A., Le, Duong T. T., Dau, Vinh Q., Peloso, Pedro L. V., Nguyen, Truong Q., Hoang, Huy D., Nguyen, Tao T. & Ziegler, Thomas, 2016, Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam, pp. 63-102 in Zootaxa 4085 (1) on pages 93-96, DOI: 10.11646/zootaxa.4085.1.3, http://zenodo.org/record/1052459, {"references":["Dubois, A. (1983) Note preliminaire sur le genre Leptolalax Dubois, 1980 (Amphibiens, Anoures), avec diagnose d'une espece novelle du Vietnam. Alytes, 2, 147 - 153.","Lathrop, A., Murphy, R. W., Orlov, N. & Ho, C. T. (1998) Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Amphibia-Reptilia, 19, 253 - 267. http: // dx. doi. org / 10.1163 / 156853898 X 00160"]}

Leptolalax ardens sp. nov. Figs. 7D, 9A, 10. Holotype. VNMN 04707/AMS R 176454, adult male, calling on clay bank 1.5 m from headwaters of 1���2m wide rocky, medium gradient, high���flow stream in evergreen forest in Kon Ka Kinh National Park, Gia Lai Province, Vietnam (14.2258�� N, 108.3230�� E; 1162 m elevation, Fig. 1, Fig 9E). Collected at 19:30 h on 21 August 2011 Jodi J. L. Rowley, Vinh Quang Dau, Huy Duc Hoang, Luong Thi Nguyen, Tan Thanh Le, Dinh Djung, and Y H'Jun. Paratypes. AMS R 176452���176453, two adult males (both calling), and VNMN 04708/AMS R 176455��� VNMN 04710/AMS R 176457, three adult males (not observed calling), collected on same date and locality as holotype. VNMN 04711 /AMS R 176458, adult female (gravid) and VNMN 04712 /AMS R 176459 adult male (calling), collected at 19:55 h on 23 August 2011 in evergreen forest in Kon Ka Kinh National Park, Gia Lai Province, Vietnam (14.2263�� N, 108.3345�� E; 1450 m elevation). UNS 00530/AMS R 176460���UNS 00532/ 176462, AMS R 176463, AMS R 176464, UNS 00533/176465, UNS 00534 /AMS R 176466, 176467, and NCSM 79650, 9 adult males, collected at night on 24 August 2011 in swampy area with small rocky stream in evergreen forest in Kon Ka Kinh National Park, Gia Lai Province, Vietnam (14.2197 �� N, 108.3207 �� E; 1041 m elevation). All specimens were collected by Jodi J. L. Rowley, Vinh Quang Dau, Huy Duc Hoang, Luong Thi Nguyen, Tan Thanh Le, Dinh Djung, and Y H'Jun. Etymology. The specific name ��� ardens ���, meaning burning, fiery, shining, or brilliant, is used as an adjective in reference to the bold and conspicuous coloration of this species compared to other species in the group. Diagnosis. Assigned to the genus Leptolalax on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra���axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax ardens sp. nov. is distinguished from its congeners by a combination of (1) supra��� axillary and ventrolateral glands present; (2) dark brownish red ventral surface with white speckling; (3) small SVL for the genus (21.6.1��� 24.7 mm in 15 adult males, 24.5 mm in one female); (4) toes lacking webbing and lateral fringes; (5) tibia length 44���49% of SVL in males; (6) pectoral gland 3.5���5.6% of SVL in males, (7) mostly smooth dorsum, (8) iris dark coppery brown, (9) distinct black supratympanic line present, (10) an advertisement call with 3���10 notes with the first, introductory note encompassing one���third of each call. Description of holotype. Head width equal to head length; snout slightly truncate in dorsal view and rounded in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than to eye; canthus rostralis indistinct, gently rounded; lores sloping; pupil vertical; eye diameter smaller than snout length; tympanum distinct, round, diameter smaller than that of the eye; tympanic rim elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings slit���like, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I Colour of holotype in life. Dorsal surface brown with irregular, darker brown blotch down back; dark interorbital bar with pale cream edging anteriorly; two small, pale markings between axilla. Blackish brown line along canthus rostralis through eye, and continuing below supratympanic ridge, terminating above axilla, encompassing nare and most of tympanum; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms orange; large, black blotch on posterior flank and numerous, smaller black spots on sides from groin to axilla (Fig. 10A,C). Upper arms pale coppery orange. Dark brownish ventral surface with white speckling on entire ventral surface including throat, arms and legs; ventral margin of throat bearing broken row of slightly larger white spots. Supra-axillary gland copper; femoral glands white; pectoral glands white, ventrolateral glands white. Iris brown; minute, black reticulations throughout. Colour of holotype in preservative. Dorsum relatively uniformly dark brown; dorsal surface of brachium, fingers and toes paler brown (Fig. 7D). Ventral surface of chest and thighs slightly medium brown with white speckles; throat dark brown speckled with white; and belly white speckled with brown. Macroglands creamy white. Measurements (mm). Holotype: SVL 22.0, HDL 9.0, HDW 8.9, SNT 3.4, EYE 2.9, IOD 2.4, TMP 1.6, TEY 0.8, TIB 10.5, EN 2.0, IN 2.5, NS 1.3, ML 5.7, PL 10.8, weight in life 1.1g Variation. Measurements of the type series are shown in Tables 2���3 and representative photographs of paratypes are shown in Figure 10. Some specimens (VNMN 04711/AMS R 176458��� VNMN 04712/1764589) have more clumped white speckling on the venter. UNS 00531/AMS R 176461 has a paler brown venter with barely distinct brown mottling. AMS R 176453, VNMN 04709/AMS R 176456, VNMN 04710/AMS R 176457, UNS 00532/AMS R 176462, AMS R 176463, UNS 00533/AMS R 176465, UNS 00534/AMS R 176466, have a distinctly pale line anterior to dark brown interorbital bar. The degree of black dorsolateral spots/flecks varies, pale edged marbling also visible on dorsum of some specimens, especially AMS R 176457, UNS 00532/AMS R 176462, UNS 00533/AMS R 176465. Advertisement call. Call descriptions are based on the calls of seven individuals, recorded at 21.4���24.7 ��C ambient temperature. Calls were an average of 239 ms in duration and consisted of 3���10 notes (Table 4, Fig. 5D). Calls were highly amplitude modulated, with amplitude peaking towards the end of each call. Calls contained 3���10 notes and introductory notes were present in most calls. Introductory notes were 25���41% of the call duration, contained up to 54 pulses. Non-introductory notes contained 1���3 pulses. Notes were repeated an average of 20 notes/s. The average dominant frequency was 3.3 kHz, spread more widely (~2.0���7.6 kHz) in non-introductory notes. Harmonics were weakly present at ~ 6, 9, and 12 kHz, and a fundamental frequency was not visible. To the human ear, the advertisement call of L. ardens sp. nov. is a weak squeak followed by a rasping, similar to an orthopteran. Ecology. All specimens of Leptolalax ardens sp. nov. were found in evergreen forest between 1,041���1,450 m elevation in Kon Ka Kinh National Park. Males were heard faintly calling on the forest floor, adjacent to small streams or seeps (Figure 9D). Distribution. Leptolalax ardens sp. nov. is only known from Kon Ka Kinh National Park, in Gia Lai Province, Vietnam. Comparisons. Leptolalax ardens sp. nov. differs from all other Leptolalax species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from L. aereus by ventral coloration, body size and the presence of a black supratympanic line, from L. bourreti by ventral coloration and body size, from L. botsfordi by male body size and relative body weight (0.41���0.47 versus 0.75���0.94 g /mm), from L. croceus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. eos by ventral coloration, male body size, iris coloration and lateral fringes on toes; from L. firthi by ventral coloration, male body size, presence of black supratympanic line, iris coloration, male skin texture and lateral fringes on toes; from L. fuliginosus by ventral coloration, male body size and iris coloration; from L. heteropus by ventral coloration and male body size; from L. kecil by ventral coloration, male body size, and iris coloration; from L. melanoleucus by ventral coloration, male body size, and iris coloration; from L. minimus by ventral coloration and male body size; from L. nahangensis by ventral coloration, male body size and iris coloration; from L. nyx by ventral coloration and male body size, from L pelodytoides by ventral coloration and male body size; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration; from L. solus by ventral coloration, male body size and iris coloration; from L. sungi by ventral coloration, male body size, presence of black supratympanic line, iris coloration and skin texture; from L. tuberosus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. ventripunctatus by ventral coloration, male body size, iris coloration and skin texture; and from L. zhangyangpingi by ventral coloration, male body size, presence of black supratympanic line, and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. ardens sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly larger body size (Wilcoxon post-hoc Z=3.7, pL. ardens sp. nov. differs from L. bidoupensis by having a smaller SVL (Wilcoxon post-hoc Z= -2.0, p=0.048), larger relative tibia length (Wilcoxon post-hoc Z=3.2, p=0.001), larger relative eye diameter (Wilcoxon post-hoc Z=2.1, p=0.038) and having a relatively uniform iris coloration (versus reddish in upper and silvery gold below). L. ardens sp. nov. differs from L. melicus by having a larger SVL (Wilcoxon post-hoc Z= -3.3, pL. ardens sp. nov. differs from L. pyrrhops, by having a smaller and non-overlapping body size (Wilcoxon post-hoc Z= 3.7, pL. ardens sp. nov. differs from L. pallidus sp. nov. by having a smaller SVL (Wilcoxon post-hoc Z= -3.8, pL. ardens sp. nov. differs from L. kalonensis sp. nov. by having a non-overlapping, smaller body size (Wilcoxon post-hoc Z= 4.7, pL. ardens sp. nov. differs from L. maculosus sp. nov. by having a smaller body size (Wilcoxon post-hoc Z=2.8, p=0.006). L. ardens sp. nov. differs from L. tadungensis sp. nov. by having a smaller body size (Wilcoxon post-hoc Z= 3.4, pL. ardens sp. nov. differs from all members of the L. applebyi group with the exception of L. kalonensis sp. nov. From L. applebyi the call of the new species differs by having a longer introductory note and note repetition rate. From L. bidoupensis, the call of L. ardens sp. nov. differs by having an introductory note in most calls and having a higher dominant frequency. From L. melicus, the call of L. ardens sp. nov. differs by having a lower dominant frequency. From L. pyrrhops the call of L. ardens sp. nov. appears to differ by having a higher dominant frequency. From L. pallidus sp. nov. the call of L. ardens sp. nov. differs by having a shorter call duration,a higher average number of pulses in the introductory note, a higher note repetition rate and a higher dominant frequency. From L. maculosus sp. nov. the call of L. ardens sp. nov. differs by having a shorter call duration, a higher number of notes per call, a distinct introductory note, and higher dominant frequency. From L. tadungensis sp. nov. the call of L. ardens sp. nov. differs by having a relatively long introductory note. No clear differences between the call of L. ardens sp. nov. and L. kalonensis sp. nov. could be determined, perhaps in part due to the limited sample size (n=1) for L. kalonensis sp. nov. See Table 7 and Figs. 5���6. Leptolalax ardens sp. nov. differs from all species within the L. applebyi group by>4.5% divergence at the 16S gene fragment examined. Interspecific variation in three L. ardens sp. nov. collected from ~ 2 km apart was 0%. 1 Leptolalax applebyi $. % $: $; L. applebyi L. L. melicus L. pyrrhops L. ardens L. pallidus L. kalonensis L. maculosus L. tadungensis bidoupensis : $;: $ +;: $ (;: $ ��;: $ 3;). (�� +. 3 (/)* 3)/ 3* ;: * K.).;:)��K.));: *(K ��;:)��K 3/;:*..K /;:(/K/ 3;:����3K3)/;:.��K +; ; * + *3 3 (+ (*�� 3 (. / *�� :*3)K).;: (*K (;: *.K. /;:*.)K *.;: K. *;:* 3K; * / *. ! Leptolalax 0 2 !> 1 2!:; 9 5 C H $ aereus H 0 0 $ $ + *K �� 3 6 A F bourreti H ��)K (B> " $ 2 K $ 0 $ botsfordi & 0 0 0 $ 3 *K (B 6 0 $ 2 $ croceus $ $ K / " 0 ��� $ eos > 0 0 0 $ *K. / 7 $ " $ $ 2 $ B $ 0 firthi > 0 (.K 3 $ $ 2 $ " B fuliginosis �� K)) B & $ " " 0 0 $ 0 heteropus 0: B = 2. (K (3; kecil 0 0 *3 K) + B 2 $ :; melanoleucus $ $ 0 $ ((K �� �� B 7 $ " " 0 0 0 minimus H + (K *. B = 2 nahangensis H 0 0 $ $.) �� B 6 $ " 0 0 nyx H 0 0 (/K (B = " 0 0 pelodytoides H " 0 / += B = " 0 0 platycephalus $ C + * B P 7 $ = $ 0 pluvialis 0 0 0 $ * K B $ 2 solus H 0 0 / (B " H $ 0 0 ������continued on the next page !:>; !> 1 2!:; 9 5 C H $ sungi H " $ $.�� K+ / $ 2 $ tuberosus H $ 0..K 3 + ��� $ $ ventripunctatus > 0 + +K ��) B> " $ $ $ 0 0 zhangyangpingi H 0 0 $ $.+ ��K+ + " $ " 0 0 0:; applebyi & 0 0 0 $ *3 (K:) ��; B 2 bidoupensis & 0 0 0 $ *�� +K +.: (; B & " 2 0 melicus & 0 0 0 $ *3 +K /:) /; B 2 pyrrhops & 0 0 0 $) ��K.:; B 7 $ 2 $ $ ardens & 0 0 0 $ * K. /: ��; B 0 2 K $ pallidus & 0 0 0 $. +K / /: + (; 0 5 kalonensis % " 0 + ��K) (: /.; B " 0 A 2 maculosus 0 " 0 $. K ((: + +; B A 2 K $ tadungensis & 0 0 0 $ K ��: +); B 2, Published as part of Rowley, Jodi J. L., Tran, Dao T. A., Le, Duong T. T., Dau, Vinh Q., Peloso, Pedro L. V., Nguyen, Truong Q., Hoang, Huy D., Nguyen, Tao T. & Ziegler, Thomas, 2016, Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam, pp. 63-102 in Zootaxa 4085 (1) on pages 75-86, DOI: 10.11646/zootaxa.4085.1.3, http://zenodo.org/record/1052459, {"references":["Dubois, A. (1983) Note preliminaire sur le genre Leptolalax Dubois, 1980 (Amphibiens, Anoures), avec diagnose d'une espece novelle du Vietnam. Alytes, 2, 147 - 153.","Lathrop, A., Murphy, R. W., Orlov, N. & Ho, C. T. (1998) Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Amphibia-Reptilia, 19, 253 - 267. http: // dx. doi. org / 10.1163 / 156853898 X 00160"]}

Leptolalax kalonensis sp. nov. Figs. 8B, 9B, 11. Holotype. IEBR A.2014.15, adult male, collected from rocky crevice, 0.5 m from 4���5 m wide, rocky stream in disturbed evergreen forest in Song Luy Watershed Forest, Binh Thuan Province, Vietnam (11.2748�� N, 108.2109�� E; 200 m elevation; Figs 1, 9F). Collected 5 August 2011 by Dao T. A. Tran, David Kizirian, Truong Q. Nguyen, Pedro L. V. Peloso, and Cuong T. Pham. Paratypes. AMNH A191768���9, 2 adult males, and IEBR A.2014.22, one adult female, collected on 6 August 2011; AMNH A191770, adult male collected on 11 August 2011; AMNH A191761 adult male collected on 12 August 2011; IEBR A.2014.18, adult male collected on 13 August 2011; all from disturbed evergreen forest in Song Luy Watershed Forest, Binh Thuan Province, Vietnam (11.2748�� N, 108.2109�� E; 200 m elevation). AMNH A191762, AMNH A191765, AMNH A191763, IEBR A.2014.19, AMNH A191764, IEBR A.2014.21, AMNH A191767, seven adult males, and AMNH A191766, IEBR A.2014.20, two adult females, collected on 5 August 2011; IEBR A.2014.15, adult male, collected on 12 August 2011; IEBR A.2014.16 and IEBR A.2014.17, two adult males, collected on 13 August 2011 in disturbed evergreen forest in Song Luy Watershed Forest, Binh Thuan Province, Vietnam (11.2919�� N, 108.2150�� E, 571 m elevation). Paratypes were collected by Dao T. A. Tran, David Kizirian, Truong Q. Nguyen, Pedro L. V. Peloso, and Cuong T. Pham. Etymology. Specific epithet ��� kalonensis ��� is in reference to the type locality of the species, which is in close proximity to the village previously known as Kalon. Diagnosis. Assigned to the genus Leptolalax on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra���axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax kalonensis sp. nov. is distinguished from its congeners by a combination of (1) supra���axillary and ventrolateral glands present; (2) pale brownish pink ventral surface with white speckling; (3) small/medium SVL for the genus (25.8���30.6 mm in 16 adult males, 28.9���30.6 mm in three females); (4) toes lacking webbing and lateral fringes; (5) tibia length 45���52% of SVL in males; (6) pectoral gland 2.5���7.0% of SVL in males, (7) mostly smooth dorsum, (8) iris gold, coppery orange in upper third in some specimens (9) distinct black supratympanic line present in most specimens, (10) an advertisement call with 4���5 notes including introductory note, comprising almost one-third of each call duration. Description of holotype. Head width slightly subequal to head length; snout slightly truncate in dorsal view and in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than eye; canthus rostralis indistinct, gently rounded; lores sloping; pupil vertical; eye diameter smaller than snout length; tympanum indistinct, round, diameter smaller than that of the eye; tympanic rim slightly elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings slit���like, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I Colour of holotype in life. Dorsal surface brown with slightly darker interorbital bar; faint dark W-shaped marking between axilla (Fig. 12A). Darker brown line along supratympanic ridge, terminating above axilla, encompassing most of tympanum; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms paler brown; several small dark brown spots on sides from groin to axilla. Dark brownish ventral surface with white speckling on ventral surfaces of throat, chest and belly, more sparse on legs and arms. Supra-axillary gland copper; femoral glands white; pectoral glands white, ventrolateral glands white. Iris copper in upper half, gold in lower half, black reticulations throughout. Colour of holotype in preservative. Dorsum greyish brown with darker brown, elongated, oval blotches and intraorbital blotch; dark brown/black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet. Groin on blotch (Fig. 8B). Ventral surface cream. Macroglands creamy white. Measurements (mm). Holotype: SVL 27.5, HDL 10.2, HDW 10.1, SNT 4.3, EYE 3.6, IOD 2.9, TMP 1.7, TEY 1.3, TIB 12.6, EN 2.5, IN 2.8, NS 1.6, ML 6.7, PL 11.5. Variation. Measurements of the type series are shown in Tables 2���3 and representative photographs of paratypes are shown in Figs. 9B and 12. AMNH A191768, AMNH A191769, IEBR A.2014.16, IEBR A.2014.17, and IEBR A.2014.22 are darker in dorsal colouration and have less distinct patterning. IEBR A.2014.18 and AMNH A191763 are pale and have only dark brown, rather than black markings. Specimens vary in degree of patterning on dorsum. Specimens vary in degree of medium brown speckling on venter; AMNH A191768, IEBR A.2014.16, AMNH A191762, IEBR A.2014.20, and AMNH A191766 have variably medium brown throats with white speckling and medium brown speckling on chest and belly (especially IEBR A.2014.16). The ventrolateral line varies in distinctness and completeness; the most distinct and complete in AMNH A191769. Advertisement call. Call descriptions are based on the call of AMNH A191770, recorded at 26.4 ��C ambient temperature. Calls were an average of 201 ms in duration and consisted of 4���5 notes (Table 7, Fig. 6B). Introductory notes of low relative amplitude were present in all calls, were an average of 29% of the call duration, and contained up to an average of 15 pulses. Non-introductory notes near-invariably contained a single pulse. Notes were repeated an average of 16 notes/s. The average dominant frequency was 2.8 kHz, spread more widely (2.6���2.9 kHz) in non-introductory notes. Harmonics and a fundamental frequency were not visible due to poor recording quality. To the human ear, the advertisement call of L. kalonensis sp. nov. is a short rasp, similar to an orthopteran. Ecology. All specimens of Leptolalax kalonensis sp. nov. were found in evergreen forest between 200���791 m elevation in Song Luy Watershed Forest. Specimens were located on the ground, 0.5���4.0 m from 2���5 m wide, shallow, rocky streams (Fig. 9F). Forest in the area was disturbed secondary forest, consisting of discontinuous patches of small hardwoods, bamboo and shrubs. Forest on mountain slopes was replaced by plantations and cultivated fields. Distribution. Leptolalax kalonensis sp. nov. is only known from Song Luy Watershed Forest in Binh Thuan Province, Vietnam. The greatest distance between known localities for the species is 43 km. Given the relatively large distance between localities, low elevation, and relatively larger streams, this species is likely to be more widely distributed than other species in the L. applebyi group. Comparisons. Leptolalax kalonensis sp. nov. differs from all other Leptolalax species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from L. aereus by ventral coloration and presence of black supratympanic line, from L. bourreti by ventral coloration, from L. botsfordi by ventral coloration, from L. croceus by ventral coloration, presence of black supratympanic line, and skin texture; from L. eos by ventral coloration, male body size, presence of black supratympanic line, and lateral fringes on toes; from L. firthi by ventral coloration, presence of black supratympanic line, skin texture and lateral fringes on toes; from L. fuliginosus by ventral coloration and iris coloration; from L. heteropus by ventral coloration and male body size; from L. kecil by ventral coloration, male body size and iris coloration; from L. melanoleucus by ventral coloration and iris coloration; from L. minimus by ventral coloration; from L. nahangensis by ventral coloration and male body size; from L. nyx by ventral coloration, from L pelodytoides by ventral coloration; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration and male body size; from L. solus by ventral coloration and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, iris coloration and skin texture; from L. ventripunctatus by ventral coloration and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. kalonensis sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly larger body size (Wilcoxon post-hoc Z=4.0, pL. kalonensis sp. nov. differs from L. bidoupensis by having a significantly larger body size (Wilcoxon post-hoc Z=4.4, pL. kalonensis sp. nov. differs from L. melicus by having a significantly larger body size (Wilcoxon post-hoc Z= -3.9, pL. kalonensis sp. nov. differs from L. pyrrhops by having a significantly smaller body size (Wilcoxon post-hoc Z=3.7, pL. kalonensis sp. nov. differs from L. ardens sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z=4.7, pL. kalonensis sp. nov. differs from L. pallidus sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z=3.0, p=0.002), smaller relative tympanum diameter (Wilcoxon post-hoc Z= -3.1, p=0.002), black supratympanic line in most specimens (versus no black supratypmanic line in L. pallidus sp. nov) and smooth skin texture (versus tuberculate in L. kalonensis sp. nov). L. kalonensis sp. nov. differs from L. maculosus sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z= -2.7, p=0.007). L. kalonensis sp. nov. differs from L. tadungensis sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z= -3.3, p=0.001), and greater relative tibia length (Wilcoxon post-hoc Z=-3.5, p=0.001). See Table 4 and Fig. 4. Although based on a single call recording, the male advertisement call of L. kalonensis sp. nov. differs from all members of the L. applebyi group with the exception of L. ardens sp. nov. In particular, the species differs from L. applebyi in length of introductory note and dominant frequency; from L. bidoupensis in call duration, presence of a distinct introductory note and dominant frequency (although similar); from L. melicus in dominant frequency; from L. pyrrhops by dominant frequency; from L. pallidus sp. nov. by average call duration, average number of pulses in the introductory note and average note repetition rate; from L. maculosus sp. nov. by average call duration, number of notes per call, and having a distinct introductory note; from L. tadungensis sp. nov. by average call duration, and average duration of introductory note. See Table 7 and Figs. 5���6. Leptolalax kalonensis sp. nov. differs from all species within the L. applebyi group by>5.2% divergence at the 16S gene fragment examined. Interspecific variation in four L. kalonensis sp. nov. collected from up to 43 km apart was 0.0���0.6%., Published as part of Rowley, Jodi J. L., Tran, Dao T. A., Le, Duong T. T., Dau, Vinh Q., Peloso, Pedro L. V., Nguyen, Truong Q., Hoang, Huy D., Nguyen, Tao T. & Ziegler, Thomas, 2016, Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam, pp. 63-102 in Zootaxa 4085 (1) on pages 90-93, DOI: 10.11646/zootaxa.4085.1.3, http://zenodo.org/record/1052459, {"references":["Dubois, A. (1983) Note preliminaire sur le genre Leptolalax Dubois, 1980 (Amphibiens, Anoures), avec diagnose d'une espece novelle du Vietnam. Alytes, 2, 147 - 153.","Lathrop, A., Murphy, R. W., Orlov, N. & Ho, C. T. (1998) Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Amphibia-Reptilia, 19, 253 - 267. http: // dx. doi. org / 10.1163 / 156853898 X 00160"]}

Leptolalax tadungensis sp. nov. Figs. 8D, 14. Holotype. UNS 00516, adult male, in dry stream in evergreen forest in Ta Dung Nature Reserve, Dak Nong Province, Vietnam (11.8350�� N, 108.0081�� E, 1262 m elevation, Fig. 1, 9G). Collected on 13 July 2010 by Dao T. A. Tran, Vinh Q. Dau, and Huy D. Hoang. Paratypes. AMS R 177666 and ZFMK 96601, two adult males, collected on same date and locality as holotype. UNS00515, AMS R 177661, AMS R 177662, ZFMK 96602 and ZFMK 96603, five adult males collected on 17 July 2010 in Ta Dung Nature Reserve, Dak Nong Province, Vietnam (11.8388�� N, 107.8945�� E, 720 m elevation). UNS 0 0 526 and UNS 0 0 527, two adult males, collected on 17 July 2011 in Ta Dung Nature Reserve, Dak Nong Province, Vietnam (11.8561�� N, 108.0371�� E, 1932 m elevation); UNS 0 0 517 adult female, collected at same site on 16 July 2011. Specimens were collected by Dao T. A. Tran and Huy D. Hoang. Etymology. Specific epithet ��� tadungensis ��� is in reference to the type locality of the species, Ta Dung Nature Reserve. Diagnosis. Assigned to the genus Leptolalax on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra���axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax tadungensis sp. nov. is distinguished from its congeners by a combination of (1) supra���axillary and ventrolateral glands present; (2) dark brownish red ventral surface with white speckling; (3) small/medium SVL for the genus (23.5���26.0 mm in eight adult males, 32.1 mm in one female); (4) toes lacking webbing and lateral fringes; (5) tibia length 43���48% of SVL in males; (6) pectoral gland 3.3���5.4% of SVL in males, (7) smooth dorsum, (8) iris dark gold, (9) distinct black supratympanic line present, (10) an advertisement call with 4���7 notes with the first, introductory note encompassing less than a tenth of each call. Description of holotype. Head width less than head length; snout rounded in dorsal view and slightly truncate in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than eye; canthus rostralis indistinct, gently rounded; lores straight; vertical pupil; eye diameter smaller than snout length; tympanum distinct, round, diameter smaller than that of the eye; tympanic rim slightly elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings slit���like, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I Colour of holotype in life. Dorsal surface dark brown; very faint transverse darker brown bars on the dorsal surface of the thighs, tibia, tarsus, lower arms, fingers and toes. Black line along supratympanic ridge, terminating above axilla, encompassing posterodorsal half of tympanum; anteroventral half paler brown; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms slightly paler, brownish copper; numerous, small darker brown spots on sides from groin to axilla. Dark brown ventral surface with fine white speckling on ventral surfaces of throat, chest, belly, legs and arms; most dense on chest. Supraaxillary gland copper; femoral glands white; pectoral glands white. Iris copper, fine black reticulations throughout. Colour of holotype in preservative. Dorsum uniform dark brown with scattered small black blotches on flanks and larger black spot on groin; fine black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet (Fig. 8D). Ventral surface medium brown with only very faintly visible, fine white speckling. Macroglands white. Measurements (mm). Holotype: SVL 24.9, HDL 9.4, HDW 8.9, SNT 3.8, EYE 2.9, IOD 3.4, TMP 1.6, TEY 0.9, TIB 11.7, EN 2.2, IN 2.2, NS 1.7, ML 6.0, PL 10.4. Variation. Measurements of the type series are shown in Tables 5���6 and representative photographs of paratypes are shown in Fig. 14. Little variation in dorsal colouration. Single female specimen UNS 0 0 527 is paler in colour with a more truncate snout. The white speckling on venter is more distinct in UNS 0 0 527, UNS 00517, and AMS R 177661, forming small blotches in UNS 0 0 527, and UNS 00517; the dorsolateral line is more distinct and less broken in these specimens. UNS 0 0 526 and UNS 00517 have a slightly shagreened skin texture in preservative, possibly an artefact of preservation (with these specimens being better preserved). Advertisement call. Call descriptions are based on the calls of four individuals, recorded at 12.9���22.3��C ambient temperature. Calls were an average of 291 ms in duration and consisted of 4���7 notes (Table 7, Figure 6D). Introductory notes were very short and quite similar to non-introductory notes, comprising only 3���9% of the call duration, and were indistinctly pulsed. Non-introductory notes contained 1���4 pulses. Notes were repeated an average of 14.3 notes/s. The average dominant frequency was 2.8 kHz. Harmonics were weakly present at ~6.3, 13.2, 16.6 kHz, and a fundamental frequency was not visible. To the human ear, the advertisement call of L. pallidus sp. nov. is a short raspy chirp, similar to an orthopteran. Ecology. All specimens of Leptolalax tadungensis sp. nov. were found in evergreen forest in Ta Dung Nature Reserve between 720���1932 m elevation (Fig. 9H). All frogs were found calling during rainy conditions. Frogs collected at the type locality were in underground rocky crevices. Individuals collected on Ta Dung Mountain (~ 1930 m elevation) were calling from the ground or under leaf litter. Distribution. Leptolalax tadungensis sp. nov. is only known from Ta Dung Nature Reserve, Dak Nong Province, Vietnam. The greatest distance between known localities for the species is 15.7 km. Comparisons. Leptolalax tadungensis sp. nov. differs from all other Leptolalax species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from L. aereus by ventral coloration and presence of distinct black supratympanic line, from L. bourreti by ventral coloration, from L. botsfordi by male body size, from L. croceus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. eos by ventral coloration, male body size, presence of black supratympanic line and lateral fringes on toes; from L. firthi by ventral coloration, iris coloration, skin texture and lateral fringes on toes; from L. fuliginosus by ventral coloration and iris coloration; from L. heteropus by ventral coloration and iris coloration; from L. kecil by ventral coloration, male body size and iris coloration; from L. melanoleucus by ventral coloration and iris coloration; from L. minimus by ventral coloration; from L. nahangensis by ventral coloration and male body size; from L. nyx by ventral coloration; from L pelodytoides by ventral coloration; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration and male body size; from L. solus by ventral coloration and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, presence of black supratympanic line and skin texture; from L. ventripunctatus by ventral coloration and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. tadungensis sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly greater body size (Wilcoxon post-hoc Z=3.6, p=0.001), greater relative diameter of the eye (Wilcoxon post-hoc Z=2.5, p=0.013), and greater relative pectoral gland size (Wilcoxon posthoc Z=2.9, p=0.004). L. tadungensis sp. nov. differs from L. bidoupensis by having a significantly smaller relative head width (Wilcoxon post-hoc Z=-2.1, p=0.032), smaller relative distance between eye and tympanum (Wilcoxon post-hoc Z=-2.1, p=0.032), and gold iris (versus reddish upper and silver below in L. bidoupensis). L. tadungensis sp. nov. differs from L. melicus by having a significantly larger body size (Wilcoxon post-hoc Z=3.5, p=0.001), and smaller relative tibia length (Wilcoxon post-hoc Z= -2.3, p=0.019). L. tadungensis sp. nov. differs from L. pyrrhops by having a significantly smaller body size (Wilcoxon post-hoc Z= -3.4, p=0.001), smaller relative tibia length (Wilcoxon post-hoc Z= -3.2, p=0.002), smaller relative distance between eye and tympanum (Wilcoxon post-hoc Z=-3.1, p=0.002) and smaller relative diameter of the eye (Wilcoxon post-hoc Z= -3.3, p=0.001). L. tadungensis sp. nov. differs from L. ardens sp. nov. by having a significantly larger body size (Wilcoxon post-hoc Z= 3.4, p=0.001) and smaller relative tibia length (Wilcoxon post-hoc Z= -3.0, p=0.003). L. tadungensis sp. nov. differs from L. pallidus sp. nov. by having smaller relative tibia length (Wilcoxon post-hoc Z= -3.1, p=0.002), smaller relative distance between eye and tympanum (Wilcoxon post-hoc Z= -2.4, p=0.019), a black supratympanic line (versus no black supratypmanic line in L. pallidus sp. nov) and mostly smooth skin texture (versus tuberculate in L. pallidus sp. nov). L. tadungensis sp. nov. differs from L. kalonensis sp. nov. by having a significantly smaller body size (Wilcoxon post-hoc Z= -3.3, p=0.001) and smaller relative tibia length (Wilcoxon post-hoc Z= -3.4, p=0.001). L. tadungensis sp. nov. differs from L. maculosus sp. nov. by having a significantly smaller relative tibia length (Wilcoxon post-hoc Z= -2.5, p=0.013). See Table 4 and Fig. 4. The male advertisement call of L. tadungensis sp. nov. differs from all species in the L. applebyi group. The call of the new species differs from L. applebyi in dominant frequency; from L. bidoupensis in having an introductory note present and in dominant frequency; from L. melicus in relative length of introductory note and dominant frequency; from L. pyrrhops by dominant frequency; from L. ardens sp. nov. by relative length of introductory note; from from L. pallidus sp. nov. by call duration and relative length of introductory note; from L. kalonensis sp. nov. by relative length of introductory note; from L. maculosus sp. nov. by call duration, number of notes per call, distinctness of introductory note and different relative duration of introductory note. See Table 7 and Fig. 5���6. Leptolalax tadungensis sp. nov. differs from all species within the L. applebyi group by>6.3% divergence at the 16S gene fragment examined. Interspecific variation in four L. tadungensis sp. nov. collected from up to ~ 15 km apart was 0.0���0.2%., Published as part of Rowley, Jodi J. L., Tran, Dao T. A., Le, Duong T. T., Dau, Vinh Q., Peloso, Pedro L. V., Nguyen, Truong Q., Hoang, Huy D., Nguyen, Tao T. & Ziegler, Thomas, 2016, Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam, pp. 63-102 in Zootaxa 4085 (1) on pages 96-99, DOI: 10.11646/zootaxa.4085.1.3, http://zenodo.org/record/1052459, {"references":["Dubois, A. (1983) Note preliminaire sur le genre Leptolalax Dubois, 1980 (Amphibiens, Anoures), avec diagnose d'une espece novelle du Vietnam. Alytes, 2, 147 - 153.","Lathrop, A., Murphy, R. W., Orlov, N. & Ho, C. T. (1998) Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Amphibia-Reptilia, 19, 253 - 267. http: // dx. doi. org / 10.1163 / 156853898 X 00160"]}

Leptolalax ardens sp. nov. Figs. 7D, 9A, 10. Holotype. VNMN 04707/AMS R 176454, adult male, calling on clay bank 1.5 m from headwaters of 1–2m wide rocky, medium gradient, high–flow stream in evergreen forest in Kon Ka Kinh National Park, Gia Lai Province, Vietnam (14.2258º N, 108.3230º E; 1162 m elevation, Fig. 1, Fig 9E). Collected at 19:30 h on 21 August 2011 Jodi J. L. Rowley, Vinh Quang Dau, Huy Duc Hoang, Luong Thi Nguyen, Tan Thanh Le, Dinh Djung, and Y H'Jun. Paratypes. AMS R 176452–176453, two adult males (both calling), and VNMN 04708/AMS R 176455– VNMN 04710/AMS R 176457, three adult males (not observed calling), collected on same date and locality as holotype. VNMN 04711 /AMS R 176458, adult female (gravid) and VNMN 04712 /AMS R 176459 adult male (calling), collected at 19:55 h on 23 August 2011 in evergreen forest in Kon Ka Kinh National Park, Gia Lai Province, Vietnam (14.2263º N, 108.3345º E; 1450 m elevation). UNS 00530/AMS R 176460–UNS 00532/ 176462, AMS R 176463, AMS R 176464, UNS 00533/176465, UNS 00534 /AMS R 176466, 176467, and NCSM 79650, 9 adult males, collected at night on 24 August 2011 in swampy area with small rocky stream in evergreen forest in Kon Ka Kinh National Park, Gia Lai Province, Vietnam (14.2197 º N, 108.3207 º E; 1041 m elevation). All specimens were collected by Jodi J. L. Rowley, Vinh Quang Dau, Huy Duc Hoang, Luong Thi Nguyen, Tan Thanh Le, Dinh Djung, and Y H'Jun. Etymology. The specific name “ ardens ”, meaning burning, fiery, shining, or brilliant, is used as an adjective in reference to the bold and conspicuous coloration of this species compared to other species in the group. Diagnosis. Assigned to the genus Leptolalax on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra–axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax ardens sp. nov. is distinguished from its congeners by a combination of (1) supra– axillary and ventrolateral glands present; (2) dark brownish red ventral surface with white speckling; (3) small SVL for the genus (21.6.1– 24.7 mm in 15 adult males, 24.5 mm in one female); (4) toes lacking webbing and lateral fringes; (5) tibia length 44–49% of SVL in males; (6) pectoral gland 3.5–5.6% of SVL in males, (7) mostly smooth dorsum, (8) iris dark coppery brown, (9) distinct black supratympanic line present, (10) an advertisement call with 3–10 notes with the first, introductory note encompassing one–third of each call. Description of holotype. Head width equal to head length; snout slightly truncate in dorsal view and rounded in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than to eye; canthus rostralis indistinct, gently rounded; lores sloping; pupil vertical; eye diameter smaller than snout length; tympanum distinct, round, diameter smaller than that of the eye; tympanic rim elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings slit–like, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I Colour of holotype in life. Dorsal surface brown with irregular, darker brown blotch down back; dark interorbital bar with pale cream edging anteriorly; two small, pale markings between axilla. Blackish brown line along canthus rostralis through eye, and continuing below supratympanic ridge, terminating above axilla, encompassing nare and most of tympanum; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms orange; large, black blotch on posterior flank and numerous, smaller black spots on sides from groin to axilla (Fig. 10A,C). Upper arms pale coppery orange. Dark brownish ventral surface with white speckling on entire ventral surface including throat, arms and legs; ventral margin of throat bearing broken row of slightly larger white spots. Supra-axillary gland copper; femoral glands white; pectoral glands white, ventrolateral glands white. Iris brown; minute, black reticulations throughout. Colour of holotype in preservative. Dorsum relatively uniformly dark brown; dorsal surface of brachium, fingers and toes paler brown (Fig. 7D). Ventral surface of chest and thighs slightly medium brown with white speckles; throat dark brown speckled with white; and belly white speckled with brown. Macroglands creamy white. Measurements (mm). Holotype: SVL 22.0, HDL 9.0, HDW 8.9, SNT 3.4, EYE 2.9, IOD 2.4, TMP 1.6, TEY 0.8, TIB 10.5, EN 2.0, IN 2.5, NS 1.3, ML 5.7, PL 10.8, weight in life 1.1g Variation. Measurements of the type series are shown in Tables 2–3 and representative photographs of paratypes are shown in Figure 10. Some specimens (VNMN 04711/AMS R 176458– VNMN 04712/1764589) have more clumped white speckling on the venter. UNS 00531/AMS R 176461 has a paler brown venter with barely distinct brown mottling. AMS R 176453, VNMN 04709/AMS R 176456, VNMN 04710/AMS R 176457, UNS 00532/AMS R 176462, AMS R 176463, UNS 00533/AMS R 176465, UNS 00534/AMS R 176466, have a distinctly pale line anterior to dark brown interorbital bar. The degree of black dorsolateral spots/flecks varies, pale edged marbling also visible on dorsum of some specimens, especially AMS R 176457, UNS 00532/AMS R 176462, UNS 00533/AMS R 176465. Advertisement call. Call descriptions are based on the calls of seven individuals, recorded at 21.4–24.7 ºC ambient temperature. Calls were an average of 239 ms in duration and consisted of 3–10 notes (Table 4, Fig. 5D). Calls were highly amplitude modulated, with amplitude peaking towards the end of each call. Calls contained 3–10 notes and introductory notes were present in most calls. Introductory notes were 25–41% of the call duration, contained up to 54 pulses. Non-introductory notes contained 1–3 pulses. Notes were repeated an average of 20 notes/s. The average dominant frequency was 3.3 kHz, spread more widely (~2.0–7.6 kHz) in non-introductory notes. Harmonics were weakly present at ~ 6, 9, and 12 kHz, and a fundamental frequency was not visible. To the human ear, the advertisement call of L. ardens sp. nov. is a weak squeak followed by a rasping, similar to an orthopteran. Ecology. All specimens of Leptolalax ardens sp. nov. were found in evergreen forest between 1,041–1,450 m elevation in Kon Ka Kinh National Park. Males were heard faintly calling on the forest floor, adjacent to small streams or seeps (Figure 9D). Distribution. Leptolalax ardens sp. nov. is only known from Kon Ka Kinh National Park, in Gia Lai Province, Vietnam. Comparisons. Leptolalax ardens sp. nov. differs from all other Leptolalax species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from L. aereus by ventral coloration, body size and the presence of a black supratympanic line, from L. bourreti by ventral coloration and body size, from L. botsfordi by male body size and relative body weight (0.41–0.47 versus 0.75–0.94 g /mm), from L. croceus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. eos by ventral coloration, male body size, iris coloration and lateral fringes on toes; from L. firthi by ventral coloration, male body size, presence of black supratympanic line, iris coloration, male skin texture and lateral fringes on toes; from L. fuliginosus by ventral coloration, male body size and iris coloration; from L. heteropus by ventral coloration and male body size; from L. kecil by ventral coloration, male body size, and iris coloration; from L. melanoleucus by ventral coloration, male body size, and iris coloration; from L. minimus by ventral coloration and male body size; from L. nahangensis by ventral coloration, male body size and iris coloration; from L. nyx by ventral coloration and male body size, from L pelodytoides by ventral coloration and male body size; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration; from L. solus by ventral coloration, male body size and iris coloration; from L. sungi by ventral coloration, male body size, presence of black supratympanic line, iris coloration and skin texture; from L. tuberosus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. ventripunctatus by ventral coloration, male body size, iris coloration and skin texture; and from L. zhangyangpingi by ventral coloration, male body size, presence of black supratympanic line, and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. ardens sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly larger body size (Wilcoxon post-hoc Z=3.7, pL. ardens sp. nov. differs from L. bidoupensis by having a smaller SVL (Wilcoxon post-hoc Z= -2.0, p=0.048), larger relative tibia length (Wilcoxon post-hoc Z=3.2, p=0.001), larger relative eye diameter (Wilcoxon post-hoc Z=2.1, p=0.038) and having a relatively uniform iris coloration (versus reddish in upper and silvery gold below). L. ardens sp. nov. differs from L. melicus by having a larger SVL (Wilcoxon post-hoc Z= -3.3, pL. ardens sp. nov. differs from L. pyrrhops, by having a smaller and non-overlapping body size (Wilcoxon post-hoc Z= 3.7, pL. ardens sp. nov. differs from L. pallidus sp. nov. by having a smaller SVL (Wilcoxon post-hoc Z= -3.8, pL. ardens sp. nov. differs from L. kalonensis sp. nov. by having a non-overlapping, smaller body size (Wilcoxon post-hoc Z= 4.7, pL. ardens sp. nov. differs from L. maculosus sp. nov. by having a smaller body size (Wilcoxon post-hoc Z=2.8, p=0.006). L. ardens sp. nov. differs from L. tadungensis sp. nov. by having a smaller body size (Wilcoxon post-hoc Z= 3.4, p The male advertisement call of L. ardens sp. nov. differs from all members of the L. applebyi group with the exception of L. kalonensis sp. nov. From L. applebyi the call of the new species differs by having a longer introductory note and note repetition rate. From L. bidoupensis, the call of L. ardens sp. nov. differs by having an introductory note in most calls and having a higher dominant frequency. From L. melicus, the call of L. ardens sp. nov. differs by having a lower dominant frequency. From L. pyrrhops the call of L. ardens sp. nov. appears to differ by having a higher dominant frequency. From L. pallidus sp. nov. the call of L. ardens sp. nov. differs by having a shorter call duration,a higher average number of pulses in the introductory note, a higher note repetition rate and a higher dominant frequency. From L. maculosus sp. nov. the call of L. ardens sp. nov. differs by having a shorter call duration, a higher number of notes per call, a distinct introductory note, and higher dominant frequency. From L. tadungensis sp. nov. the call of L. ardens sp. nov. differs by having a relatively long introductory note. No clear differences between the call of L. ardens sp. nov. and L. kalonensis sp. nov. could be determined, perhaps in part due to the limited sample size (n=1) for L. kalonensis sp. nov. See Table 7 and Figs. 5–6. Leptolalax ardens sp. nov. differs from all species within the L. applebyi group by>4.5% divergence at the 16S gene fragment examined. Interspecific variation in three L. ardens sp. nov. collected from ~ 2 km apart was 0%. 1 Leptolalax applebyi $. % $: $; L. applebyi L. L. melicus L. pyrrhops L. ardens L. pallidus L. kalonensis L. maculosus L. tadungensis bidoupensis : $;: $ +;: $ (;: $ ­;: $ 3;). (­ +. 3 (/)* 3)/ 3* ;: * K.).;:)­K.));: *(K ­;:)­K 3/;:*..K /;:(/K/ 3;:­­3K3)/;:.­K +; : K(;:+K­;:.K/;:+K(;: K*);:.K/;:.K+;:/K ­;:.K/; % % % % % % 9 %:;:; M *: K3; *:*K; N*/ + 3 * (:.K.;: +K.*;: K (;:*)K /;: K3; K K / /:+K*; *.­ *+.(K:*(K /;:)K+.;: /K(;:* K­­; K *:*K+; * *:*K; * * *:*K; * * * *:*K.; * /:*K.; ** *(*­ *­ (*3 ­ (­ *(*(+ *. :­ 3K*.;:*. (K*­;:*+ 3K * *;:*(­K*3.;:*+ *K 3 3;:(K­);:** *K;:* ­K*3 /; . * * / *: *K.; + ­ /: /K ­; ­ : ’A;: 3K.;:* 3K;: /K ­;:* 3K;:.K /;: (K *;:O>; * + *3 3 (+ (*­ 3 (. / *­ :*3)K).;: (*K (;: *.K. /;:*.)K *.;: K. *;:* 3K; * / *. ! Leptolalax 0 2 !> 1 2!:; 9 5 C H $ aereus H 0 0 $ $ + *K ­ 3 6 A F bourreti H ­)K (B> " $ 2 K $ 0 $ botsfordi & 0 0 0 $ 3 *K (B 6 0 $ 2 $ croceus $ $ K / " 0 ’ $ eos > 0 0 0 $ *K. / 7 $ " $ $ 2 $ B $ 0 firthi > 0 (.K 3 $ $ 2 $ " B fuliginosis ­ K)) B & $ " " 0 0 $ 0 heteropus 0: B = 2. (K (3; kecil 0 0 *3 K) + B 2 $ :; melanoleucus $ $ 0 $ ((K ­ ­ B 7 $ " " 0 0 0 minimus H + (K *. B = 2 nahangensis H 0 0 $ $.) ­ B 6 $ " 0 0 nyx H 0 0 (/K (B = " 0 0 pelodytoides H " 0 / += B = " 0 0 platycephalus $ C + * B P 7 $ = $ 0 pluvialis 0 0 0 $ * K B $ 2 solus H 0 0 / (B " H $ 0 0 ……continued on the next page !:>; !> 1 2!:; 9 5 C H $ sungi H " $ $.­ K+ / $ 2 $ tuberosus H $ 0..K 3 + ’ $ $ ventripunctatus > 0 + +K ­) B> " $ $ $ 0 0 zhangyangpingi H 0 0 $ $.+ ­K+ + " $ " 0 0 0:; applebyi & 0 0 0 $ *3 (K:) ­; B 2 bidoupensis & 0 0 0 $ *­ +K +.: (; B & " 2 0 melicus & 0 0 0 $ *3 +K /:) /; B 2 pyrrhops & 0 0 0 $) ­K.:; B 7 $ 2 $ $ ardens & 0 0 0 $ * K. /: ­; B 0 2 K $ pallidus & 0 0 0 $. +K / /: + (; 0 5 kalonensis % " 0 + ­K) (: /.; B " 0 A 2 maculosus 0 " 0 $. K ((: + +; B A 2 K $ tadungensis & 0 0 0 $ K ­: +); B 2

Leptolalax pallidus sp. nov. Figs. 8A, 11. Holotype. UNS 0 0 510, adult male, calling on plant leaf 0.1 m from ground, 1 m from 2���3 m wide rocky stream in evergreen forest, Bidoup Nui Ba National Park, Lam Dong Province, Vietnam. (12.1520 �� N, 108.695�� E; 1644 m elevation, Fig. 1, 9E). Collected at night on 27 June 2011 by Dao T. A. Tran and Duong T. T. Le. Paratypes. UNS 0 0 512, AMS R 177657, ZFMK 96599, three adult males collected on 26 July 2011 in in evergreen forest in Bidoup Nui Ba National Park, Lam Dong Province, Vietnam (12.1519�� N, 108.6940�� E; 1681 m elevation). UNS 0 0 511, AMS R 177659, ZFMK 96598, three adult males collected on same date and location as holotype. AMS R 177658, collected on 27 June 2011 at same location as holotype. All specimens were collected by Dao T. A. Tran and Duong T. T. Le Etymology. The specific name ��� pallidus ���, meaning pale, is used as an adjective in reference to the paler coloration and less distinct markings of this species, particularly in preservative, compared to other species in the group. Diagnosis. Assigned to the genus Leptolalax on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra���axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax pallidus sp. nov. is distinguished from its congeners by a combination of (1) supra���axillary and ventrolateral glands present; (2) dark brownish red ventral surface with faint white speckling; (3) small SVL for the genus (24.5.1��� 27.7 mm in 8 adult males), (4) toes lacking webbing and lateral fringes; (5) tibia length 45���51% of SVL in males; (6) pectoral gland 3.4���6.7% of SVL in males, (7) tuberculate skin texture on dorsum, (8) iris copper in upper half, gold in lower half; (9) distinct black supratympanic line absent, (10) an advertisement call with 4���7 notes with the short introductory note encompassing a quarter of each call and only one note in non-introductory notes. Description of holotype. Head width slightly subequal to head length; snout truncate in dorsal view and in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than to eye; canthus rostralis indistinct, gently rounded; lores sloping; pupil vertical; eye diameter smaller than snout length; tympanum indistinct, round, diameter smaller than that of the eye; tympanic rim slightly elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings small, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I Colour of holotype in life. Dorsal surface brown with slightly darker interorbital bar with pale brown patches anteriorly and posteriorly; faint dark W-shaped marking between axilla (Fig. 11A). Darker brown line along supratympanic ridge, terminating above axilla, encompassing most of tympanum; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms paler brown; numerous, small dark brown spots on sides from groin to axilla. Dark brownish ventral surface with white speckling on ventral surfaces of throat, chest and belly, more sparse on legs and arms. Supra-axillary gland copper; femoral glands white; pectoral glands white, ventrolateral glands white. Iris copper in upper half, gold in lower half, black reticulations throughout. Colour of holotype in preservative. Dorsum relatively uniform medium brown with darker interorbital bar and slightly paler posterior to this; dorsal surface of brachium, fingers and toes paler brown, dark brown/black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet (Fig. 8A). Elbows and upper arms pale brown. Ventral surface medium brown with faint white speckles. Macroglands creamy white. Measurements (mm). Holotype: SVL 25.2, HDL 9.4, HDW 9.2, SNT 3.9, EYE 3.4, IOD 3.2, TMP 1.7, TEY 1.4, TIB 13.0, EN 2.5, IN 2.6, NS 1.2, ML 6.8, PL 12.6. Variation. Measurements of the type series are shown in Tables 5���6 and representative photographs of paratypes are shown in Fig. 11. All specimens have a degree of darker brown patterning (mostly marbling but also some spots) on the dorsum and dark banding on the tibiotarsus and antebrachium, hands and feet, but both are more pronounced in UNS00512, AMS R 177657 and UNS00511. ZFMK 96599 has faint dark brown spots evenly distributed around tubercles on dorsum. UNS00510���00512 have more distinctly pale elbows in preservative. Size and distribution of white speckles on the venter varies a little between specimens, more clumped on chest in AMS R 177659 and ZFMK 96598. Skin texture less tuberculate in preservative than in life. Advertisement call. Call descriptions are based on the calls of three individuals, recorded at 14.0���21.4 ��C ambient temperature. Calls were an average of 673 ms in duration and consisted of 4���7 notes (Table 7, Fig. 6A). Short introductory notes of low relative amplitude were present in all calls. Introductory notes were 22���26% of the call duration, contained up to 63 pulses. Non-introductory notes invariably contained a single pulse. Amplitude peaked towards the middle of the call. Notes were repeated an average of 6.8 notes/s. The average dominant frequency was 2.5 kHz. Harmonics were weakly present at ~7.3 kHz, and a fundamental frequency was not visible. To the human ear, the advertisement call of L. pallidus sp. nov. consists of a squelch followed by clicking. Ecology. All specimens of Leptolalax pallidus sp. nov. were found in evergreen forest in Bidoup Nui Ba National Park between 1644���1681 m elevation. Males were located on the forest floor, on leaf-litter or rocky crevices and clay holes in the banks of a dry stream. Fig 9E. Distribution. Leptolalax pallidus sp. nov. is only known from Gia Rich mountain in Bidoup Nui Ba National Park, only about 5 km from L. bidoupensis. The two species occur in continuous forest, with no obvious current biogeographic barrier between them. Comparisons. Leptolalax pallidus sp. nov. differs from all other Leptolalax species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from L. aereus by ventral coloration and iris coloration, from L. bourreti by ventral coloration, male body size, absence of black supratympanic line, skin texture and iris coloration, from L. botsfordi by male body size, absence of black supratympanic line, iris coloration and relative male body weight (0.42���0.50 versus 0.75���0.94 g /mm), from L. croceus by ventral coloration and iris coloration; from L. eos by ventral coloration, male body size, iris coloration and lateral fringes on toes; from L. firthi by ventral coloration, absence of black supratympanic line, iris coloration and lateral fringes on toes; from L. fuliginosus by ventral coloration, male body size, skin texture and iris coloration; from L. heteropus by ventral coloration, male body size, skin texture and absence of black supratympanic line; from L. kecil by ventral coloration, male body size, absence of black supratympanic line and iris coloration; from L. melanoleucus by ventral coloration, body size, absence of black supratympanic line, skin texture and iris coloration; from L. minimus by ventral coloration, skin texture and male body size; from L. nahangensis by ventral coloration, male body size, skin texture, absence of black supratympanic line and iris coloration; from L. nyx by ventral coloration, skin texture and absence of black supratympanic line, from L pelodytoides by ventral coloration and absence of black supratympanic line; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration, body size, skin texture and absence of black supratympanic line; from L. solus by ventral coloration, absence of black supratympanic line, skin texture and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. ventripunctatus by ventral coloration, absence of black supratympanic line, skin texture and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size, skin texture and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. pallidus sp. nov. differs by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly larger and non-overlapping male body size (Wilcoxon post-hoc Z=3.4, pL. applebyi) and tuberculate skin texture (versus mostly smooth in L. applebyi). L. pallidus sp. nov. differs from L. bidoupensis by having a significantly larger body size (Wilcoxon post-hoc Z=2.7, p=0.006), larger relative tibia length (Wilcoxon post-hoc Z=3.0, p=0.002), larger relative tympanum size (Wilcoxon post-hoc Z=2.0, p=0.041), dark brown iris (versus reddish upper and silver below), indistinct, brown supratympanic line (versus distinct black supratypmanic line in L. bidoupensis) and tuberculate skin texture (versus mostly smooth in L. bidoupensis). L. pallidus sp. nov. differs from L. melicus by having a significantly larger and non-overlapping male body size (Wilcoxon post-hoc Z=-3.3, pL. applebyi) and tuberculate skin texture (versus mostly smooth in L. melicus). L. pallidus sp. nov. differs from L. pyrrhops by having a significantly smaller and non-overlapping male body size (Wilcoxon post-hoc Z=3.2, p=0.002), smaller relative diameter of the eye (Wilcoxon post-hoc Z=3.2, p=0.002), indistinct, brown supratympanic line (versus distinct black supratypmanic line in L. pyrrhops) and tuberculate skin texture (versus only slightly shagreened in L. pyrrhops). L. pallidus sp. nov. differs from L. ardens sp. nov. by having a significantly larger body size (Wilcoxon post-hoc Z=-3.8, pL. ardens sp. nov) and tuberculate skin texture (versus mostly smooth in L. ardens sp. nov). L. pallidus sp. nov. differs from L. kalonensis sp. nov. by having a significantly smaller body size (Wilcoxon post-hoc Z=3.0, p=0.002), larger relative tympanum diameter (Wilcoxon post-hoc Z=-3.1, p=0.002), indistinct, brown supratympanic line (versus distinct black supratypmanic line in most L. kalonensis sp. nov) and tuberculate skin texture (versus mostly smooth in L. kalonensis sp. nov). L. pallidus sp. nov. differs from L. maculosus sp. nov. by having indistinct, brown supratympanic line (versus distinct black supratypmanic line in L. maculosus sp. nov) and tuberculate skin texture (versus mostly smooth in L. maculosus sp. nov). L. pallidus sp. nov. differs from L. tadungensis sp. nov. by having a significantly larger relative tibia length (Wilcoxon post-hoc Z=-3.1, p=0.002), larger relative distance between the eye and tympanum (Wilcoxon post-hoc Z=-2.4, p=0.019), indistinct, brown supratympanic line (versus distinct black supratympanic line in L. maculosus sp. nov) and tuberculate skin texture (versus mostly smooth in L. maculosus sp. nov). See Table 4 and Fig. 4. The male advertisement call of L. pallidus sp. nov. differs from all members of the L. applebyi group. The call of L. pallidus sp. nov. differs from L. applebyi in call duration, having a relatively long, pulsed introductory note, having one pulse in non-introductory notes, and by average note repetition rate and average dominant frequency. L. pallidus sp. nov. differs from L. bidoupensis in call duration, in having a distinct, pulsed introductory note, and by average note repetition rate and perhaps average dominant frequency (non-overlapping but similar values); from L. melicus in call duration, number of pulses in the introductory note, average note repetition rate and average dominant frequency; from L. pyrrhops in call duration, average number of pulses in introductory note, average note repetition rate and potentially by dominant frequency; from L. ardens sp. nov. differs by call duration, number of pulses in the introductory note, note repetition rate and dominant frequency; from L. kalonensis sp. nov. in average call duration, average number of pulses in the introductory note and average note repetition rate; from L. maculosus sp. nov. in average call duration number of notes per call, and having a distinct introductory note; from L. tadungensis sp. nov. in average call duration, average duration of introductory note, and note repetition rate. See Table 7 and Figs. 5���6. Leptolalax pallidus sp. nov. differs from all species within the L. applebyi group by>4.7% divergence at the 16S gene fragment examined. Interspecific variation in three L. pallidus sp. nov. collected from, Published as part of Rowley, Jodi J. L., Tran, Dao T. A., Le, Duong T. T., Dau, Vinh Q., Peloso, Pedro L. V., Nguyen, Truong Q., Hoang, Huy D., Nguyen, Tao T. & Ziegler, Thomas, 2016, Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam, pp. 63-102 in Zootaxa 4085 (1) on pages 87-90, DOI: 10.11646/zootaxa.4085.1.3, http://zenodo.org/record/1052459, {"references":["Dubois, A. (1983) Note preliminaire sur le genre Leptolalax Dubois, 1980 (Amphibiens, Anoures), avec diagnose d'une espece novelle du Vietnam. Alytes, 2, 147 - 153.","Lathrop, A., Murphy, R. W., Orlov, N. & Ho, C. T. (1998) Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Amphibia-Reptilia, 19, 253 - 267. http: // dx. doi. org / 10.1163 / 156853898 X 00160"]}

Leptolalax kalonensis sp. nov. Figs. 8B, 9B, 11. Holotype. IEBR A.2014.15, adult male, collected from rocky crevice, 0.5 m from 4–5 m wide, rocky stream in disturbed evergreen forest in Song Luy Watershed Forest, Binh Thuan Province, Vietnam (11.2748º N, 108.2109º E; 200 m elevation; Figs 1, 9F). Collected 5 August 2011 by Dao T. A. Tran, David Kizirian, Truong Q. Nguyen, Pedro L. V. Peloso, and Cuong T. Pham. Paratypes. AMNH A191768–9, 2 adult males, and IEBR A.2014.22, one adult female, collected on 6 August 2011; AMNH A191770, adult male collected on 11 August 2011; AMNH A191761 adult male collected on 12 August 2011; IEBR A.2014.18, adult male collected on 13 August 2011; all from disturbed evergreen forest in Song Luy Watershed Forest, Binh Thuan Province, Vietnam (11.2748º N, 108.2109º E; 200 m elevation). AMNH A191762, AMNH A191765, AMNH A191763, IEBR A.2014.19, AMNH A191764, IEBR A.2014.21, AMNH A191767, seven adult males, and AMNH A191766, IEBR A.2014.20, two adult females, collected on 5 August 2011; IEBR A.2014.15, adult male, collected on 12 August 2011; IEBR A.2014.16 and IEBR A.2014.17, two adult males, collected on 13 August 2011 in disturbed evergreen forest in Song Luy Watershed Forest, Binh Thuan Province, Vietnam (11.2919º N, 108.2150º E, 571 m elevation). Paratypes were collected by Dao T. A. Tran, David Kizirian, Truong Q. Nguyen, Pedro L. V. Peloso, and Cuong T. Pham. Etymology. Specific epithet “ kalonensis ” is in reference to the type locality of the species, which is in close proximity to the village previously known as Kalon. Diagnosis. Assigned to the genus Leptolalax on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra–axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax kalonensis sp. nov. is distinguished from its congeners by a combination of (1) supra–axillary and ventrolateral glands present; (2) pale brownish pink ventral surface with white speckling; (3) small/medium SVL for the genus (25.8–30.6 mm in 16 adult males, 28.9–30.6 mm in three females); (4) toes lacking webbing and lateral fringes; (5) tibia length 45–52% of SVL in males; (6) pectoral gland 2.5–7.0% of SVL in males, (7) mostly smooth dorsum, (8) iris gold, coppery orange in upper third in some specimens (9) distinct black supratympanic line present in most specimens, (10) an advertisement call with 4–5 notes including introductory note, comprising almost one-third of each call duration. Description of holotype. Head width slightly subequal to head length; snout slightly truncate in dorsal view and in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than eye; canthus rostralis indistinct, gently rounded; lores sloping; pupil vertical; eye diameter smaller than snout length; tympanum indistinct, round, diameter smaller than that of the eye; tympanic rim slightly elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings slit–like, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I Colour of holotype in life. Dorsal surface brown with slightly darker interorbital bar; faint dark W-shaped marking between axilla (Fig. 12A). Darker brown line along supratympanic ridge, terminating above axilla, encompassing most of tympanum; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms paler brown; several small dark brown spots on sides from groin to axilla. Dark brownish ventral surface with white speckling on ventral surfaces of throat, chest and belly, more sparse on legs and arms. Supra-axillary gland copper; femoral glands white; pectoral glands white, ventrolateral glands white. Iris copper in upper half, gold in lower half, black reticulations throughout. Colour of holotype in preservative. Dorsum greyish brown with darker brown, elongated, oval blotches and intraorbital blotch; dark brown/black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet. Groin on blotch (Fig. 8B). Ventral surface cream. Macroglands creamy white. Measurements (mm). Holotype: SVL 27.5, HDL 10.2, HDW 10.1, SNT 4.3, EYE 3.6, IOD 2.9, TMP 1.7, TEY 1.3, TIB 12.6, EN 2.5, IN 2.8, NS 1.6, ML 6.7, PL 11.5. Variation. Measurements of the type series are shown in Tables 2–3 and representative photographs of paratypes are shown in Figs. 9B and 12. AMNH A191768, AMNH A191769, IEBR A.2014.16, IEBR A.2014.17, and IEBR A.2014.22 are darker in dorsal colouration and have less distinct patterning. IEBR A.2014.18 and AMNH A191763 are pale and have only dark brown, rather than black markings. Specimens vary in degree of patterning on dorsum. Specimens vary in degree of medium brown speckling on venter; AMNH A191768, IEBR A.2014.16, AMNH A191762, IEBR A.2014.20, and AMNH A191766 have variably medium brown throats with white speckling and medium brown speckling on chest and belly (especially IEBR A.2014.16). The ventrolateral line varies in distinctness and completeness; the most distinct and complete in AMNH A191769. Advertisement call. Call descriptions are based on the call of AMNH A191770, recorded at 26.4 ºC ambient temperature. Calls were an average of 201 ms in duration and consisted of 4–5 notes (Table 7, Fig. 6B). Introductory notes of low relative amplitude were present in all calls, were an average of 29% of the call duration, and contained up to an average of 15 pulses. Non-introductory notes near-invariably contained a single pulse. Notes were repeated an average of 16 notes/s. The average dominant frequency was 2.8 kHz, spread more widely (2.6–2.9 kHz) in non-introductory notes. Harmonics and a fundamental frequency were not visible due to poor recording quality. To the human ear, the advertisement call of L. kalonensis sp. nov. is a short rasp, similar to an orthopteran. Ecology. All specimens of Leptolalax kalonensis sp. nov. were found in evergreen forest between 200–791 m elevation in Song Luy Watershed Forest. Specimens were located on the ground, 0.5–4.0 m from 2–5 m wide, shallow, rocky streams (Fig. 9F). Forest in the area was disturbed secondary forest, consisting of discontinuous patches of small hardwoods, bamboo and shrubs. Forest on mountain slopes was replaced by plantations and cultivated fields. Distribution. Leptolalax kalonensis sp. nov. is only known from Song Luy Watershed Forest in Binh Thuan Province, Vietnam. The greatest distance between known localities for the species is 43 km. Given the relatively large distance between localities, low elevation, and relatively larger streams, this species is likely to be more widely distributed than other species in the L. applebyi group. Comparisons. Leptolalax kalonensis sp. nov. differs from all other Leptolalax species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from L. aereus by ventral coloration and presence of black supratympanic line, from L. bourreti by ventral coloration, from L. botsfordi by ventral coloration, from L. croceus by ventral coloration, presence of black supratympanic line, and skin texture; from L. eos by ventral coloration, male body size, presence of black supratympanic line, and lateral fringes on toes; from L. firthi by ventral coloration, presence of black supratympanic line, skin texture and lateral fringes on toes; from L. fuliginosus by ventral coloration and iris coloration; from L. heteropus by ventral coloration and male body size; from L. kecil by ventral coloration, male body size and iris coloration; from L. melanoleucus by ventral coloration and iris coloration; from L. minimus by ventral coloration; from L. nahangensis by ventral coloration and male body size; from L. nyx by ventral coloration, from L pelodytoides by ventral coloration; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration and male body size; from L. solus by ventral coloration and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, iris coloration and skin texture; from L. ventripunctatus by ventral coloration and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. kalonensis sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly larger body size (Wilcoxon post-hoc Z=4.0, pL. kalonensis sp. nov. differs from L. bidoupensis by having a significantly larger body size (Wilcoxon post-hoc Z=4.4, pL. kalonensis sp. nov. differs from L. melicus by having a significantly larger body size (Wilcoxon post-hoc Z= -3.9, pL. kalonensis sp. nov. differs from L. pyrrhops by having a significantly smaller body size (Wilcoxon post-hoc Z=3.7, pL. kalonensis sp. nov. differs from L. ardens sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z=4.7, pL. kalonensis sp. nov. differs from L. pallidus sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z=3.0, p=0.002), smaller relative tympanum diameter (Wilcoxon post-hoc Z= -3.1, p=0.002), black supratympanic line in most specimens (versus no black supratypmanic line in L. pallidus sp. nov) and smooth skin texture (versus tuberculate in L. kalonensis sp. nov). L. kalonensis sp. nov. differs from L. maculosus sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z= -2.7, p=0.007). L. kalonensis sp. nov. differs from L. tadungensis sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z= -3.3, p=0.001), and greater relative tibia length (Wilcoxon post-hoc Z=-3.5, p=0.001). See Table 4 and Fig. 4. Although based on a single call recording, the male advertisement call of L. kalonensis sp. nov. differs from all members of the L. applebyi group with the exception of L. ardens sp. nov. In particular, the species differs from L. applebyi in length of introductory note and dominant frequency; from L. bidoupensis in call duration, presence of a distinct introductory note and dominant frequency (although similar); from L. melicus in dominant frequency; from L. pyrrhops by dominant frequency; from L. pallidus sp. nov. by average call duration, average number of pulses in the introductory note and average note repetition rate; from L. maculosus sp. nov. by average call duration, number of notes per call, and having a distinct introductory note; from L. tadungensis sp. nov. by average call duration, and average duration of introductory note. See Table 7 and Figs. 5–6. Leptolalax kalonensis sp. nov. differs from all species within the L. applebyi group by>5.2% divergence at the 16S gene fragment examined. Interspecific variation in four L. kalonensis sp. nov. collected from up to 43 km apart was 0.0–0.6%.

Theloderma lateriticum sp. nov. Figs. 2, 3 Holotype: AMNH 168757 / IEBR A. 0860, adult male, Vietnam, Lao Cai Province, Van Ban District, Nam Tha Commune, above the Nam Tha river in disturbed, submontane semi-evergreen forest near 21 ° 54 ' 56 " N, 104 ° 21 ' 39 " E, between 1300–1400 m elevation, 10 September 2004, Raoul H. Bain, Nguyen Quang Truong, Lu A Cho & Trieu Tai Vuong. The holotype has an incision in the left leg from tissue samplings of muscle. Diagnosis: Theloderma lateriticum sp. nov. is distinguishable from all other rhacophorid species from South China, Northeast India, and mainland Southeast Asia (north of the Isthmus of Kra) by a combination of: male SVL 23.5 mm; tympanum distinct, with distinct tympanic annulus; vomerine teeth absent; hand webbing absent; foot webbing reduced (Toe I, below proximal subarticular tubercle on postaxial side; Toe II, to level of proximal subarticular tubercle on postaxial side; Toe III below proximal subarticular tubercle on preaxial side, and just beyond proximal subarticular tubercle on postaxial side; Toe IV, to level of proximal subarticular tubercle on preaxial side, and just beyond proximal subarticular tubercle on postaxial side; and Toe V, just beyond proximal subarticular tubercle on preaxial side); outer metatarsal tubercle present; dorsal skin granular, bearing keratinized spicules, raised on small, isolated bumps; dermal fringes on the post axial portions of the limbs absent; dorsum, iris brick-red; mid-dorsal spot black; lip with small white spots; nuptial pad on Finger I; vocal sacs absent. Description of holotype: AMNH 168757 / IEBR A. 0860, adult male, habitus slender; head narrow, longer than wide; snout subacuminate in dorsal view, rounded in lateral aspect, projecting beyond lower jaw; nostril lateral, nearer to tip of snout than eye; canthus rostralis distinct, but not raised, distinct sulcus between nares; lores slightly concave, almost vertical, oblique; eye diameter 67 % of snout length; pupil horizontally oval; interorbital distance greater than width of upper eyelid; pineal body not visible; tympanum distinct, round, covered by layer of skin, 74 % of eye diameter, not depressed relative to skin of temporal region, tympanic annulus thin; vomerine teeth absent; choanae at extreme lateral edge; tongue elongated-cordiform, deeply notched posteriorly, free for approximately two-thirds its length; vocal sacs absent. Tips of all four fingers expanded into round discs; circummarginal and transverse grooves appear as one continuous circular groove; width of Finger I disc smallest, width of Finger III disc about 1.5 times width of phalanx, 67 % diameter of tympanum; relative Finger lengths I Tips of toes expanded into rounded discs, smaller than width of fingers; circummarginal and transverse grooves appear as one continuous circular groove; relative Toe lengths I Dorsal skin granular, bearing keratinized spicules on small, isolated bumps; keratinized spicules on small, isolated bumps also present on posterior portion of thigh around cloaca and preaxial surfaces of limb, but absent from lips, loreal region, and tympanum; supratympanic fold indistinct, curves from posterior edge of eye anterior to arm insertion, thickening considerably posterior to tympanum; rictal glands absent; postaxial line of tubercles on the limbs saw-tooth-like, not pronounced; tarsal fold absent; throat smooth, becoming coarsely tubercular towards belly. Measurements (mm) of holotype: SVL 23.5; HDL 9.5; HDW 7.9; SNT 3.8; IND: 2.3; EYE 2.6; IOD 2.8; TMP 1.9; TEY 0.5; HND 7.2; F 3 DSC: 1.3; F 4 DSC 1.0; TIB 12.7; FEM 10.6; FTL 9.6. Color in life: Dorsum, snout, loreal region, upper lip, upper eyelids, supratympanic fold, and all accompanying tubercles with deep brick-red wash overtop dark black-green; lips with small white spots; black mid-dorsal spot; dorsal wash fades to black-green patch on the sacral area; iris deep brick-red; tympanum black with bright white dots in the middle; upper portion of flanks deep brick-red with black blotches, including one large blotch in communication with tympanum and the largest in the sacro-inguinal region; lower half of flank with bright white spots interspersed among row-like black blotches; limbs barred black on red-brown to hands, feet; digit tips black with white spots; tubercles on limbs bright white to digit tips; venter uniform grey-brown with bright white spots, most dense on chest and belly. Color in preservative: Dorsum, snout, loreal region, upper lip, upper eyelids, supratympanic fold, with light grey sheen over light brown spot on mid portion of head and dark grey elsewhere; lips, tympanum redbrown with dull white spotting; black mid-dorsal spot clearly visible as in life; flank light brown with black blotches clearly visible as in life; sacral area dark grey, with two dark dorsolateral black lines; limbs barred black with red-brown; digit tips grey; anterior portion of thigh mottled light brown with light grey; posterior portion of thigh light grey with light brown dusting; all dorsal tubercles white; throat, ventral portions of limbs as in color in life; chest with white spots around dark brown curved lines; each ‘grain’ of granular belly uniquely colored white brown or grey to form a patchwork; ventral portion of hands light grey; ventromarginal grooves of toes light grey. Habitat and ecology: This specimen was collected from inside a bamboo hollow from a disturbed, submontane, evergreen forest. The locality is close to a cardamom farm within a bamboo forest that is inundated with small streams and seeps. The temperature had reached a daytime high of 31 o C, and a nighttime low of 19 o C. It was breezy. Also present inside bamboo of the same area were specimens of Theloderma asperum (adults and egg clutches), Kurixalus odontotarsus, and the snake Rhadinophis prasinus. Because of its locality in far northwestern Vietnam, this species may also occur in neighboring southeastern Yunnan Province, China, and northeastern Laos. Etymology: The Latin adjective lateriticum means ‘of bricks’, referring to the brick-red color of the dorsum Comparisons: The generic assignment of many small rhacophorid frog species from Asia is still uncertain (e.g. Bossuyt & Dubois 2001; Wilkinson et al. 2002; Frost et al. 2006; Grismer et al. 2007; Li et al. 2008; Fei et al., 2009). For this reason, we compare the new species with all species of Theloderma, and all rhacophorids from Northeast India, South China (including Hainan Island), and mainland Southeast Asia (north of the Isthmus of Kra) whose adult males have SVL less than 35 mm, lack hand webbing, lack dermal fringes on the post axial portions of the limbs, lack green coloration, and lack dorsal stripes: i.e., Theloderma asperum; T. bicolor; T. corticale; T. gordoni; T. horridum; T. kwangsiense; T. leporosum; T. licin; T. moloch; T. nagalandense; T. phrynoderma; T. rhododiscus; T. ryabovi; T. stellatum; Chiromantis laevis; Nyctixalus pictus; Philautus abditus; P. andersoni; P. aurifasciatus; P. cardamonus; P. cinerascens; P. garo; P. hainanus; P. jinxiuensis; P. k e m p i a e; P. k e m p i i; P. longchuanensis; P. maosonensis; P. m e n g l a e n s i s; P. ocellatus; P. parvulus; P. petilus; P. shillongensis; P. truongsonensis; P. tytthus. Theloderma lateriticum sp. nov. can be immediately distinguished from all known species of Theloderma by its brick-red dorsal wash (T. asperum, T. rhododiscus, and T. ryabovi dominated by black and white; T. bicolor, T. corticale, and T. kwangsiense dominated by green and brown; T. gordoni, T. horridum, T. leporosum dominated by brown; T. licin solid white in the day, mottled with brown at night and in captivity; T. nagalandense dominated by large, orange-red patches; T. moloch, T. phrynoderma dominated by grey; and T. stellatum dominated by brown or lavender-grey), and minimal foot webbing: Toe I, below proximal subarticular tubercle on postaxial side (versus at least to subarticular tubercle); Toe II, to level of proximal subarticular tubercle on postaxial side (versus at least beyond subarticular tubercle); Toe III below proximal subarticular tubercle on preaxial side, and just beyond proximal subarticular tubercle on postaxial side (versus at least beyond subarticular tubercle on preaxial side, and at least to distal subarticular tubercle on postaxial side); Toe IV, to level of proximal subarticular tubercle on preaxial side, and just beyond proximal subarticular tubercle on postaxial side (versus at least to medial subarticular tubercle on preaxial side of IV, and at least to distal subarticular tubercle on postaxial side); and Toe V, just beyond proximal subarticular tubercle on preaxial side (versus at least to distal subarticular tubercle). Theloderma lateriticum sp. nov. (SVL 23.5 mm) can further be distinguished from large congeners T. bicolor, T. corticale, T. gordoni, T. horridum, T. kwangsiense, T. leporosa, T. moloch, T. nagalandense, T. phrynoderma, and T. ryabovi (adult male SVL 40 mm or greater), by its distinct tympanic annulus (absent in all large congeners), and its granular dorsum (heavily rugose in all large congeners, except T. ryabovi). In addition, T. ryabovi and T. horridum males possess vocal sacs (absent in T. lateriticum). . There are four small species of Theloderma (adult male SVL 29–35 mm, T. asperum; 28, 30 mm, T. licin; 26, 27 mm, T. rhododiscus; and 27–35 mm, T. stellatum). Theloderma lateriticum sp. nov. further differs from T. asperum with its line of tubercles on post axial portions of the limb (absent in T. asperum), and absence of vocal sacs in males (present in males of T. asperum). Theloderma lateriticum sp. nov. further differs from T. rhododiscus by having a granular dorsum with small, isolated bumps (rough, with network of longitudinal pointy ridges in T. rhododiscus), one black mid-dorsal spot (several large dorsal spots in T. rhododiscus), and grey ventral digital surfaces (orange-red in T. rhododiscus). Theloderma lateriticum sp. nov. further differs from T. stellatum with its free hands (fingers one-third webbed in T. stellatum), finger discs smaller than tympanum diameter (at least as large as tympanum in T. stellatum), snout spotted white (spotted black in T. stellatum). Theloderma lateriticum sp. nov. most closely resembles the brown form of T. licin, but in addition to the above, further differs in the absence of finger webbing (present in T. licin), the presence of a nuptial pad on Finger I (present on Finger II in T. licin), the presence of tubercles on the supratympanic fold (absent in T. licin), and the absence of vocal sacs in males (present in males of T. licin). The brick-red dorsum also differentiates Theloderma lateriticum sp. nov. from other regional rhacophorids that have male SVL less than 35 mm, lack hand webbing, lack dermal fringes on the post axial portions of the limbs, lack green coloration, and lack dorsal stripes: white with red vermiculations in C. laevis; dark and/or cinnamon brown in N. pictus; olive, grey, or brown in P. abditus, P. andersoni, P. cardamonus, P. cinerascens, P. g a ro, P. j i n x i u e n s i s, P. k e m p i a e, P. k e m p i i, P. longchuanensis, P. maosonensis, P. menglaensis, P. ocellatus, P. parvulus, P. shillongensis, P. tytthus; dorsolateral portions soft yellow-beige in P. petilus; yellow-gold with dark brown dorsolateral edges in P. truongsonensis; and variable but never red in P. aurifasciatus. Theloderma lateriticum sp. nov. can be further differentiated from these rhacophorids by a combination of the following: snout length longer than eye diameter (opposite condition in N. pictus, P. abditus, P. cinerascens, P. garo, P. kempiae, P. parvulus); absence of vomerine teeth (present in N. pictus, P. petilus); absence of vocal sacs (present in Philautus abditus, P. aurifasciatus, P. cardamonus, P. hainanus, P. maosonensis, P. menglaensis; P. ocellatus, P. parvulus, P. petilus); distinct tympanum (completely hidden in P. abditus, P. kempiae, P. m a o s o n e n s i s; partially obscured in P. aurifasciatus, P. parvulus, P. tytthus), an outer metatarsal tubercle (absent in C. laevis, P. abditus, P. cardamonus, P. hainanus, P. k e m p i i, P. longchuanensis, and P. t y t t h u s); upper surfaces with keratinized spicules (absent in C. laevis, Philautus abditus, P. cardamonus, P. g a ro, P. jinxiuensis, P. longchuanensis, P. truongsonensis); postaxial limb tubercles (absent in P. petilus, N. pictus), and solid dorsal wash (with interorbital sash, dorsal butterfly, ‘)(’, or X marking in Philautus abditus, P. andersoni, P. cardamonus, P. c i n e r a s c e n s, P. garo, P. jinxiuensis, P. k e m p i a e, P. maosonensis, P. menglaensis, P. parvulus, P. shillongensis, P. tytthus; snout a different shade than rest of dorsum in P. kempii, P. shillongensis). Theloderma lateriticum sp. nov. further differs from N. pictus by its supratympanic fold (absent in N. pictus); banded limbs (unbanded in N. pictus); and venter with spots (immaculate in N. pictus). Theloderma lateriticum sp. nov. further differs from P. andersoni by its reduced foot webbing (for T. lateriticum sp. nov. web below proximal subarticular tubercle on Toe I, preaxial sides of II, III [at or beyond level of proximal subarticular tubercle in P. andersoni]; web to level of proximal subarticular tubercle on postaxial side of Toe II, preaxial side of IV, and just beyond proximal subarticular tubercle on postaxial side of Toe III, IV, preaxial side of V [well beyond level of proximal subarticular tubercle in P. andersoni]); its banded limbs (not banded in P. andersoni); and its grey-brown venter (yellow in P. andersoni). Theloderma lateriticum sp. nov. further differs from P. aurifasciatus by its reduced foot webbing (for T. lateriticum sp. nov. web below proximal subarticular tubercle on Toe I, preaxial side of II [at level of proximal subarticular tubercle in P. aurifasciatus]; web to level of proximal subarticular tubercle on preaxial side of IV, and just beyond proximal subarticular tubercle on postaxial side of Toe III, IV, preaxial side of V [well beyond level of proximal subarticular tubercle in P. aurifasciatus]). Theloderma lateriticum sp. nov. further differs from P. k e m p i i in having a disc on FI (absent in P. k e m p i i), and an inner metatarsal tubercle (absent in P. k e m p i i). Theloderma lateriticum sp. nov. further differs from P. ocellatus by its brick-red iris (yellow in P. ocellatus) and spotted lip (banded in P. ocellatus). Theloderma lateriticum sp. nov. further differs from P. petilus with its minimal foot webbing (to discs as a fringe on every toe in P. petilus), tubercular belly, and spinose cloacal tubercles on posterior portion of thigh (both areas smooth in P. petilus). Theloderma lateriticum sp. nov. further differs from P. truongsonensis by its long head (HDW = HDL in P. truongsonensis), finger subarticular tubercle formula of 1,1,2,2 (1,2,2,1 in P. truongsonensis), Toe III being shorter than Toe V (III = V in P. truongsonensis), and lack of tarsal fold (present in P. truongsonensis). Theloderma lateriticum sp. nov. further differs from P. shillongensis by its long head (HDW> HDL in P. shillongensis), pedal web (only basal web between Toes IV, and V, otherwise free in P. shillongensis), and Toe III being shorter than Toe V (opposite condition in P. shillongensis). Remarks: McLeod & Ahmad (2007) noted that T. licin has the ability to change color. They further suggested that the color of a preserved specimen may be related to its color at the time of euthanasia. In our field experience, the Vietnam populations of the following species do not change color: T. asperum, T. bicolor, T. corticale, T. gordoni, T. rhododiscus, and T. stellatum.

Theloderma lateriticum sp. nov. Figs. 2, 3 Holotype: AMNH 168757 / IEBR A. 0860, adult male, Vietnam, Lao Cai Province, Van Ban District, Nam Tha Commune, above the Nam Tha river in disturbed, submontane semi-evergreen forest near 21 �� 54 ' 56 " N, 104 �� 21 ' 39 " E, between 1300���1400 m elevation, 10 September 2004, Raoul H. Bain, Nguyen Quang Truong, Lu A Cho & Trieu Tai Vuong. The holotype has an incision in the left leg from tissue samplings of muscle. Diagnosis: Theloderma lateriticum sp. nov. is distinguishable from all other rhacophorid species from South China, Northeast India, and mainland Southeast Asia (north of the Isthmus of Kra) by a combination of: male SVL 23.5 mm; tympanum distinct, with distinct tympanic annulus; vomerine teeth absent; hand webbing absent; foot webbing reduced (Toe I, below proximal subarticular tubercle on postaxial side; Toe II, to level of proximal subarticular tubercle on postaxial side; Toe III below proximal subarticular tubercle on preaxial side, and just beyond proximal subarticular tubercle on postaxial side; Toe IV, to level of proximal subarticular tubercle on preaxial side, and just beyond proximal subarticular tubercle on postaxial side; and Toe V, just beyond proximal subarticular tubercle on preaxial side); outer metatarsal tubercle present; dorsal skin granular, bearing keratinized spicules, raised on small, isolated bumps; dermal fringes on the post axial portions of the limbs absent; dorsum, iris brick-red; mid-dorsal spot black; lip with small white spots; nuptial pad on Finger I; vocal sacs absent. Description of holotype: AMNH 168757 / IEBR A. 0860, adult male, habitus slender; head narrow, longer than wide; snout subacuminate in dorsal view, rounded in lateral aspect, projecting beyond lower jaw; nostril lateral, nearer to tip of snout than eye; canthus rostralis distinct, but not raised, distinct sulcus between nares; lores slightly concave, almost vertical, oblique; eye diameter 67 % of snout length; pupil horizontally oval; interorbital distance greater than width of upper eyelid; pineal body not visible; tympanum distinct, round, covered by layer of skin, 74 % of eye diameter, not depressed relative to skin of temporal region, tympanic annulus thin; vomerine teeth absent; choanae at extreme lateral edge; tongue elongated-cordiform, deeply notched posteriorly, free for approximately two-thirds its length; vocal sacs absent. Tips of all four fingers expanded into round discs; circummarginal and transverse grooves appear as one continuous circular groove; width of Finger I disc smallest, width of Finger III disc about 1.5 times width of phalanx, 67 % diameter of tympanum; relative Finger lengths I Measurements (mm) of holotype: SVL 23.5; HDL 9.5; HDW 7.9; SNT 3.8; IND: 2.3; EYE 2.6; IOD 2.8; TMP 1.9; TEY 0.5; HND 7.2; F 3 DSC: 1.3; F 4 DSC 1.0; TIB 12.7; FEM 10.6; FTL 9.6. Color in life: Dorsum, snout, loreal region, upper lip, upper eyelids, supratympanic fold, and all accompanying tubercles with deep brick-red wash overtop dark black-green; lips with small white spots; black mid-dorsal spot; dorsal wash fades to black-green patch on the sacral area; iris deep brick-red; tympanum black with bright white dots in the middle; upper portion of flanks deep brick-red with black blotches, including one large blotch in communication with tympanum and the largest in the sacro-inguinal region; lower half of flank with bright white spots interspersed among row-like black blotches; limbs barred black on red-brown to hands, feet; digit tips black with white spots; tubercles on limbs bright white to digit tips; venter uniform grey-brown with bright white spots, most dense on chest and belly. Color in preservative: Dorsum, snout, loreal region, upper lip, upper eyelids, supratympanic fold, with light grey sheen over light brown spot on mid portion of head and dark grey elsewhere; lips, tympanum redbrown with dull white spotting; black mid-dorsal spot clearly visible as in life; flank light brown with black blotches clearly visible as in life; sacral area dark grey, with two dark dorsolateral black lines; limbs barred black with red-brown; digit tips grey; anterior portion of thigh mottled light brown with light grey; posterior portion of thigh light grey with light brown dusting; all dorsal tubercles white; throat, ventral portions of limbs as in color in life; chest with white spots around dark brown curved lines; each ���grain��� of granular belly uniquely colored white brown or grey to form a patchwork; ventral portion of hands light grey; ventromarginal grooves of toes light grey. Habitat and ecology: This specimen was collected from inside a bamboo hollow from a disturbed, submontane, evergreen forest. The locality is close to a cardamom farm within a bamboo forest that is inundated with small streams and seeps. The temperature had reached a daytime high of 31 o C, and a nighttime low of 19 o C. It was breezy. Also present inside bamboo of the same area were specimens of Theloderma asperum (adults and egg clutches), Kurixalus odontotarsus, and the snake Rhadinophis prasinus. Because of its locality in far northwestern Vietnam, this species may also occur in neighboring southeastern Yunnan Province, China, and northeastern Laos. Etymology: The Latin adjective lateriticum means ���of bricks���, referring to the brick-red color of the dorsum Comparisons: The generic assignment of many small rhacophorid frog species from Asia is still uncertain (e.g. Bossuyt & Dubois 2001; Wilkinson et al. 2002; Frost et al. 2006; Grismer et al. 2007; Li et al. 2008; Fei et al., 2009). For this reason, we compare the new species with all species of Theloderma, and all rhacophorids from Northeast India, South China (including Hainan Island), and mainland Southeast Asia (north of the Isthmus of Kra) whose adult males have SVL less than 35 mm, lack hand webbing, lack dermal fringes on the post axial portions of the limbs, lack green coloration, and lack dorsal stripes: i.e., Theloderma asperum; T. bicolor; T. corticale; T. gordoni; T. horridum; T. kwangsiense; T. leporosum; T. licin; T. moloch; T. nagalandense; T. phrynoderma; T. rhododiscus; T. ryabovi; T. stellatum; Chiromantis laevis; Nyctixalus pictus; Philautus abditus; P. andersoni; P. aurifasciatus; P. cardamonus; P. cinerascens; P. garo; P. hainanus; P. jinxiuensis; P. k e m p i a e; P. k e m p i i; P. longchuanensis; P. maosonensis; P. m e n g l a e n s i s; P. ocellatus; P. parvulus; P. petilus; P. shillongensis; P. truongsonensis; P. tytthus. Theloderma lateriticum sp. nov. can be immediately distinguished from all known species of Theloderma by its brick-red dorsal wash (T. asperum, T. rhododiscus, and T. ryabovi dominated by black and white; T. bicolor, T. corticale, and T. kwangsiense dominated by green and brown; T. gordoni, T. horridum, T. leporosum dominated by brown; T. licin solid white in the day, mottled with brown at night and in captivity; T. nagalandense dominated by large, orange-red patches; T. moloch, T. phrynoderma dominated by grey; and T. stellatum dominated by brown or lavender-grey), and minimal foot webbing: Toe I, below proximal subarticular tubercle on postaxial side (versus at least to subarticular tubercle); Toe II, to level of proximal subarticular tubercle on postaxial side (versus at least beyond subarticular tubercle); Toe III below proximal subarticular tubercle on preaxial side, and just beyond proximal subarticular tubercle on postaxial side (versus at least beyond subarticular tubercle on preaxial side, and at least to distal subarticular tubercle on postaxial side); Toe IV, to level of proximal subarticular tubercle on preaxial side, and just beyond proximal subarticular tubercle on postaxial side (versus at least to medial subarticular tubercle on preaxial side of IV, and at least to distal subarticular tubercle on postaxial side); and Toe V, just beyond proximal subarticular tubercle on preaxial side (versus at least to distal subarticular tubercle). Theloderma lateriticum sp. nov. (SVL 23.5 mm) can further be distinguished from large congeners T. bicolor, T. corticale, T. gordoni, T. horridum, T. kwangsiense, T. leporosa, T. moloch, T. nagalandense, T. phrynoderma, and T. ryabovi (adult male SVL 40 mm or greater), by its distinct tympanic annulus (absent in all large congeners), and its granular dorsum (heavily rugose in all large congeners, except T. ryabovi). In addition, T. ryabovi and T. horridum males possess vocal sacs (absent in T. lateriticum). . There are four small species of Theloderma (adult male SVL 29���35 mm, T. asperum; 28, 30 mm, T. licin; 26, 27 mm, T. rhododiscus; and 27���35 mm, T. stellatum). Theloderma lateriticum sp. nov. further differs from T. asperum with its line of tubercles on post axial portions of the limb (absent in T. asperum), and absence of vocal sacs in males (present in males of T. asperum). Theloderma lateriticum sp. nov. further differs from T. rhododiscus by having a granular dorsum with small, isolated bumps (rough, with network of longitudinal pointy ridges in T. rhododiscus), one black mid-dorsal spot (several large dorsal spots in T. rhododiscus), and grey ventral digital surfaces (orange-red in T. rhododiscus). Theloderma lateriticum sp. nov. further differs from T. stellatum with its free hands (fingers one-third webbed in T. stellatum), finger discs smaller than tympanum diameter (at least as large as tympanum in T. stellatum), snout spotted white (spotted black in T. stellatum). Theloderma lateriticum sp. nov. most closely resembles the brown form of T. licin, but in addition to the above, further differs in the absence of finger webbing (present in T. licin), the presence of a nuptial pad on Finger I (present on Finger II in T. licin), the presence of tubercles on the supratympanic fold (absent in T. licin), and the absence of vocal sacs in males (present in males of T. licin). The brick-red dorsum also differentiates Theloderma lateriticum sp. nov. from other regional rhacophorids that have male SVL less than 35 mm, lack hand webbing, lack dermal fringes on the post axial portions of the limbs, lack green coloration, and lack dorsal stripes: white with red vermiculations in C. laevis; dark and/or cinnamon brown in N. pictus; olive, grey, or brown in P. abditus, P. andersoni, P. cardamonus, P. cinerascens, P. g a ro, P. j i n x i u e n s i s, P. k e m p i a e, P. k e m p i i, P. longchuanensis, P. maosonensis, P. menglaensis, P. ocellatus, P. parvulus, P. shillongensis, P. tytthus; dorsolateral portions soft yellow-beige in P. petilus; yellow-gold with dark brown dorsolateral edges in P. truongsonensis; and variable but never red in P. aurifasciatus. Theloderma lateriticum sp. nov. can be further differentiated from these rhacophorids by a combination of the following: snout length longer than eye diameter (opposite condition in N. pictus, P. abditus, P. cinerascens, P. garo, P. kempiae, P. parvulus); absence of vomerine teeth (present in N. pictus, P. petilus); absence of vocal sacs (present in Philautus abditus, P. aurifasciatus, P. cardamonus, P. hainanus, P. maosonensis, P. menglaensis; P. ocellatus, P. parvulus, P. petilus); distinct tympanum (completely hidden in P. abditus, P. kempiae, P. m a o s o n e n s i s; partially obscured in P. aurifasciatus, P. parvulus, P. tytthus), an outer metatarsal tubercle (absent in C. laevis, P. abditus, P. cardamonus, P. hainanus, P. k e m p i i, P. longchuanensis, and P. t y t t h u s); upper surfaces with keratinized spicules (absent in C. laevis, Philautus abditus, P. cardamonus, P. g a ro, P. jinxiuensis, P. longchuanensis, P. truongsonensis); postaxial limb tubercles (absent in P. petilus, N. pictus), and solid dorsal wash (with interorbital sash, dorsal butterfly, ���)(���, or X marking in Philautus abditus, P. andersoni, P. cardamonus, P. c i n e r a s c e n s, P. garo, P. jinxiuensis, P. k e m p i a e, P. maosonensis, P. menglaensis, P. parvulus, P. shillongensis, P. tytthus; snout a different shade than rest of dorsum in P. kempii, P. shillongensis). Theloderma lateriticum sp. nov. further differs from N. pictus by its supratympanic fold (absent in N. pictus); banded limbs (unbanded in N. pictus); and venter with spots (immaculate in N. pictus). Theloderma lateriticum sp. nov. further differs from P. andersoni by its reduced foot webbing (for T. lateriticum sp. nov. web below proximal subarticular tubercle on Toe I, preaxial sides of II, III [at or beyond level of proximal subarticular tubercle in P. andersoni]; web to level of proximal subarticular tubercle on postaxial side of Toe II, preaxial side of IV, and just beyond proximal subarticular tubercle on postaxial side of Toe III, IV, preaxial side of V [well beyond level of proximal subarticular tubercle in P. andersoni]); its banded limbs (not banded in P. andersoni); and its grey-brown venter (yellow in P. andersoni). Theloderma lateriticum sp. nov. further differs from P. aurifasciatus by its reduced foot webbing (for T. lateriticum sp. nov. web below proximal subarticular tubercle on Toe I, preaxial side of II [at level of proximal subarticular tubercle in P. aurifasciatus]; web to level of proximal subarticular tubercle on preaxial side of IV, and just beyond proximal subarticular tubercle on postaxial side of Toe III, IV, preaxial side of V [well beyond level of proximal subarticular tubercle in P. aurifasciatus]). Theloderma lateriticum sp. nov. further differs from P. k e m p i i in having a disc on FI (absent in P. k e m p i i), and an inner metatarsal tubercle (absent in P. k e m p i i). Theloderma lateriticum sp. nov. further differs from P. ocellatus by its brick-red iris (yellow in P. ocellatus) and spotted lip (banded in P. ocellatus). Theloderma lateriticum sp. nov. further differs from P. petilus with its minimal foot webbing (to discs as a fringe on every toe in P. petilus), tubercular belly, and spinose cloacal tubercles on posterior portion of thigh (both areas smooth in P. petilus). Theloderma lateriticum sp. nov. further differs from P. truongsonensis by its long head (HDW = HDL in P. truongsonensis), finger subarticular tubercle formula of 1,1,2,2 (1,2,2,1 in P. truongsonensis), Toe III being shorter than Toe V (III = V in P. truongsonensis), and lack of tarsal fold (present in P. truongsonensis). Theloderma lateriticum sp. nov. further differs from P. shillongensis by its long head (HDW> HDL in P. shillongensis), pedal web (only basal web between Toes IV, and V, otherwise free in P. shillongensis), and Toe III being shorter than Toe V (opposite condition in P. shillongensis). Remarks: McLeod & Ahmad (2007) noted that T. licin has the ability to change color. They further suggested that the color of a preserved specimen may be related to its color at the time of euthanasia. In our field experience, the Vietnam populations of the following species do not change color: T. asperum, T. bicolor, T. corticale, T. gordoni, T. rhododiscus, and T. stellatum., Published as part of Bain, Raoul H., Nguyen, Truong Q. & Doan, Kien V., 2009, A new species of the genus Theloderma Tschudi, 1838 (Anura: Rhacophoridae) from Northwestern Vietnam, pp. 58-68 in Zootaxa 2191 on pages 60-65, DOI: 10.5281/zenodo.189385, {"references":["Bossuyt, F. & Dubois, A. (2001) A review of the frog genus Philautus Gistel, 1848 (Amphibia, Anura, Ranidae, Rhacophorinae). Zeylanica, 6, 1 - 112.","Wilkinson, J. A., Drewes, R. C. & Tatum, O. L. (2002) A molecular phylogenetic analysis of the family Rhacophoridae with an emphasis on the Asian and African genera. Molecular Phylogenetics and Evolution, 24, 265 - 273.","Frost, D. R., Grant, T., Faivovich, J., Bain, R. 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