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Dendrelaphis in Vietnam is provided below: 1a. Vertebral scale row distinctly enlarged, two supralabials touching the eye........................................ 2 1b. Vertebral scale row feebly enlarged, one supralabial touching the eye............................................ 3 2a. Ventrolateral black stripe present................................................................. D. pictus 2b. Ventrolateral black stripe absent............................................................ D. ngansonensis 3a. Dorsal scale rows 15 on midbody............................................................. D. subocularis 3b. Dorsal scale rows 13 on midbody........................................................... D. binhi sp. nov., Published as part of Nguyen, Sang Ngoc, Nguyen, Vu Dang Hoang, Le, Manh Van, Nguyen, Luan Thanh, Vo, Thi-Dieu-Hien, Vo, Ba Dinh, Che, Jing & Murphy, Robert W., 2023, A new snake of the genus Dendrelaphis Boulenger, 1890 (Squamata: Colubridae) from the coastal area of southern Vietnam, pp. 130-144 in Zootaxa 5318 (1) on page 142, DOI: 10.11646/zootaxa.5318.1.6, http://zenodo.org/record/8158335

Dendrelaphis binhi sp. nov. Holotype. ITBCZ 6663, adult male, collected from Tuy Phong, Binh Thuan Province, Vietnam; coordinates 11 o 12’36.2”N, 108 o 41’43.2”E; elevation 34 m a.s.l. by S.N. Nguyen and L. T. Nguyen, on 23 July 2018 (Figs. 3 & 4). Paratypes. Seven specimens: ITBCZ 6664, ITBCZ 6666 (adult females), ITBCZ 6665 (subadult male), collected by the same collectors, on the same date and site as the holotype; ITBCZ 6681 (adult male), collected from Tuy Phong, Binh Thuan Province, Vietnam, coordinates 11 o 13’03.3”N, 108 o 39’10.2”E, elevation 39 m a.s.l. by S.N. Nguyen and L. T. Nguyen, on 26 July 2018; ITBCZ 5944-6 (adult females), collected from Thuan Nam, Ninh Thuan Province, Vietnam; coordinates 11 o 19’18.4”N, 108 o 56’46.6”E; elevation 105 m a.s.l. by S.N. Nguyen and D.H. V. Nguyen, on 13 April 2017 (Figs. 3–5). Diagnosis. Dendrelaphis binhi sp. nov. is distinguished from all of its congeners by a combination of the following morphological characters: medium sized bronzeback snake (largest TL 936 mm in female); one long supralabial entering orbit; body scale smooth, 13 rows on neck and midbody, 9–11 before vent; vertebral scale row slightly enlarged; ventrals 154–158 in males and 161–170 in females; subcaudals 105 or 106 in males and 95–102 in females; 18 or 19 maxillary teeth, posterior teeth shorter than the others; hemipenis spinose, reaching 10 th or 11 th SC, with a tiny papilla; cloacal plate divided; dorsum bright brown with a white stripe along the lower flank. Description of holotype. Adult male; head elongate (HL/HW = 1.99), distinct from neck; body elongate, SVL 495 mm; tail long (TaL/TL = 0.305), 217 mm in length; eye quite large with round pupil, eye diameter shorter than snout length (ED/SnL: 0.76) (Table 3). Head scalation. Dorsal head scales smooth, abutting along midline rather than overlapping; rostral broader than high, visible from above, touching nasals, internasals and 1 st supralabial on both sides; supralabials 8/8, first fours small, the 5 th bordering eye on both sides, the 6 th small and triangle shaped, the 7 th and 8 th distinct larger than the 6 th and the first fours; 1/1 loreal, much longer than high (2.0 mm length vs. 0.8 mm height), touching nasal, internasal, prefrontal, preocular and the 2 nd and 3 rd supralabials; 1/1 preocular; 2/2 postoculars; 2+2/2+2 temporals; suture between internasals as long as that between prefrontals; frontal bell-shaped, widened anteriorly and much narrowed posteriorly (5.6 mm length, 3.6 mm width); parietals longer than frontal (6.5 mm vs. 5.6 mm length), bordered laterally by upper temporals and postocular and anteriorly by frontal and supraoculars; 10/10 infralabials, first pair medially in contact with each other, first five in contact with anterior chin shield, the 6 th largest and touching posterior chin shield; posterior chin shields slightly longer than anterior ones (5.1 mm vs. 4.9 mm) (Fig. 4A). Body scalation. Dorsal scales smooth, in 13-13-9 rows; lateral scales, except for the outer row, disposed obliquely; vertebral scale row slightly enlarged, narrower than the outer row of scales; scale row reductions from 13 to 11 at ventrals 89 and 90 and from 11 to 9 at ventrals 96 and 99; ventrals 158, with a suture-like lateral keel; cloacal plate divided; subcaudals 106, all paired; terminal caudal scale forming a pointed cap. Dentition. Maxillary teeth 18, continuously, the posterior teeth slightly smaller than the others. Hemipenis. Hemipenis unforked, reaching 11 th SC, spinose throughout, except for the apical naked area. The spines in the terminal half of the organ are largest. Sulcus spermaticus single, extending to tip, the sulcus lip prominent. On the apical naked area, there is a tiny lobule with small spines (Fig. 4B&C). Coloration. In life, dorsal and lateral parts of the body and tail bronze-brown, laterally this color extends to the upper half of the second scale row; upper part of head olive, lip and lower parts of the head and neck yellow; iris orange on the upper part and black on anterior, posterior, and lower parts of the eye; concealed parts of lateral scales light blue; a black temporal streak extending from eye to neck; a bright vertebral stripe on the anterior part of dorsum; two lateral stripes: a white one on the lower half of the second scale row and the upper half of the outer row and the second stripe is pale brown occurring on the lower half of the outer row and the edge of ventral scale; anterior part of the body with a series of black blotches at the suture between the lateral edge of ventral and the outer row, these blotches are less distinct or absent on the middle and posterior parts of the body; ventral white. In preservation, color faded but the pattern remained, all yellow parts becoming white. Variation. Paratypes ITBCZ 6665 and ITBCZ 5945 have 7 and 9 supralabials on the right side, respectively; paratypes ITBCZ 5945 and ITBCZ 5946 have 11 dorsal scale rows before vent; paratype ITBCZ 5946 has a short suture on the lower margin of the 5 th supralabial on both sides; paratype ITBCZ 5945 has the upper half of the second dorsal row black forming a black longitudinal line (Fig. 3E); lip and lower parts of head and neck yellow in all specimens collected in July in Tuy Phong, Binh Thuan Province but white in all snakes collected in April in Thuan Nam, Ninh Thuan Province. Table 3 summarizes variation in size and scalation of type series. Sexual dimorphism. Male has fewer ventral scales (V 154–158 in 3 males, 161–170 in 5 females) and slightly longer relative tail (TaL/TL 0.305–0.309 in 2 adult males; 0.277–0.297 in 5 adult females). Etymology. We name the new species in honor of Dr. Ngô Văn Bὶnh, a Vietnamese herpetologist who passed away suddenly in 2023 at the age of 49 by a serious stroke. We recommend “Binh’s Bronzeback Snake” and “Rắn leo cây bὶnh” as the common English and Vietnamese names of the new species, respectively. Distribution. The new species is currently known from Thuan Nam, Ninh Thuan Province and Tuy Phong, Binh Thuan Province, Vietnam (Fig. 1). Field notes. All specimens were collected at night, between 7:00 PM and 11:00 PM, while sleeping on brushes at about 1–2 m above ground. The habitat in Thuan Nam, Ninh Thuan Province is a coastal hill with brushes and granite outcrops (Fig. 5B) and the habitat in Tuy Phong, Binh Thuan Province is an agricultural area with scattered large brushes. The holotype was collected on a large brush group together with paratypes ITBCZ 6664 and ITBCZ 6665 and with two other individuals of Ahaetulla fusca (Dumeril, Bibron & Dumeril). Gravid paratype ITBCZ 5946 has three eggs. Comparisons. Dendrelaphis binhi sp. nov. differs morphologically from all congeners in Indochina as follows: from D. biloreatus (Wall, 1908) by having fewer ventrals (154–170 vs. 187–199), fewer subcaudals (95–106 vs. 139–145), one (vs. two) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Smith 1943); from D. caudolineatus by having fewer ventrals in males (154–158 vs. 171–180), fewer ventrals in females (161–170 vs. 171–185), one (vs. two) supralabial entering the eye, and absence (vs. presence) of black longitudinal stripes along the entire length of the body (Rooijen & Vogel 2012); from D. cyanochloris (Wall, 1921) by having fewer midbody scale rows (13 vs. 15), fewer ventrals (154–170 vs. 181–206), fewer subcaudals (95–106 vs. 135–157), one (vs. two) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Rooijen & Vogel 2008; Jiang et al. 2020); from D. formosus by having fewer midbody scale rows (13 vs. 15), fewer ventrals (154–170 vs. 172–194), fewer subcaudals (95–106 vs. 142–162), and one (vs. two) supralabial entering the eye (Jiang et al. 2020); from D. haasi by having fewer midbody scale rows (13 vs. 15), one (vs. two) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Das 2015); from D. kopsteini Vogel & Van Rooijen, 2007 by having fewer midbody scale rows (13 vs. 15), one (vs. two) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Vogel & Rooijen 2007); from D. ngansonensis by having fewer subcaudals (95–106 vs. 117–153), one (vs. two or three) supralabial entering the eye, vertebral scale row slightly enlarged (vs. strongly enlarged), fewer maxillary teeth (18 or 19 vs. 24 or 25), and shorter hemipenis (extending to SC 10 or 11 vs. SC 18) (Ziegler & Vogel 1999; Jiang et al. 2020); from D. nigroserratus Vogel, Rooijen & Hauser, 2012 by having fewer midbody scale rows (13 vs. 15), fewer ventrals (154–170 vs. 197– 204), fewer subcaudals (95–106 vs. 148–152), one (vs. three) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Vogel et al. 2012); from D. pictus by having fewer midbody scale rows (13 vs. 15), fewer subcaudals (95–106 vs. 113–148), fewer maxillary teeth (18 or 19 vs. 23–28), one (vs. two or three) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Smith 1943; Vogel & Rooijen 2008); from D. striatus by having fewer midbody scale rows (13 vs. 15), one (vs. two or three) supralabial entering the eye, and absence (vs. presence) of black oblique bars laterally on the body (Ziegler & Vogel 1999; Vogel & Rooijen 2007); from D. subocularis by having fewer dorsal scale rows at neck (13 vs. 15) and at midbody (13 vs. 15), and fewer maxillary teeth (18 or 19 vs. 21–23) (Smith 1943; Rooijen & Vogel 2010); from D. tristis by having fewer midbody scale rows (13 vs. 15), fewer subcaudals (95–106 vs. 108–145), one (vs. two) supralabial entering the eye, and longer hemipenis (extending to SC 10 or 11 vs. SC 8) (Smith 1943); from D. vogeli by having fewer midbody scale rows (13 vs. 15), one (vs. two) supralabial entering the eye, fewer ventrals (154–170 vs. 193–197), fewer subcaudals (95–106 vs. 130–135), and vertebral scale row slightly enlarged (vs. strongly enlarged) (Jiang et al. 2020); and from D. walli Vogel & Rooijen, 2011 by having fewer ventrals (154–170 vs. 193–197), fewer subcaudals (95–106 vs. 130–135), one (vs. two) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Vogel & Rooijen 2011)., Published as part of Nguyen, Sang Ngoc, Nguyen, Vu Dang Hoang, Le, Manh Van, Nguyen, Luan Thanh, Vo, Thi-Dieu-Hien, Vo, Ba Dinh, Che, Jing & Murphy, Robert W., 2023, A new snake of the genus Dendrelaphis Boulenger, 1890 (Squamata: Colubridae) from the coastal area of southern Vietnam, pp. 130-144 in Zootaxa 5318 (1) on pages 136-139, DOI: 10.11646/zootaxa.5318.1.6, http://zenodo.org/record/8158335, {"references":["Smith, M. A. (1943) The fauna of British India, including Ceylon and Burma: Reptilia and Amphibia. Taylor and Francis, London, 583 pp.","Rooijen, J. V. & Vogel, G. (2012) A revision of the taxonomy of Dendrelaphis caudolineatus (Gray, 1834) (Serpentes: Colubridae). Zootaxa, 3272 (1), 1 - 25. https: // doi. org / 10.11646 / zootaxa. 3272.1.1","Rooijen, J. V. & Vogel, G. (2008) A new species of Dendrelaphis (Serpentes: Colubridae) from Java, Indonesia. The Raffles Bulletin of Zoology, 56, 189 - 197.","Jiang, K., Ren, J. L., Guo, J. P., Wang, Z., Ding, L. & Li, J. T. (2020) A new species of the genus Dendrelaphis (Squamata: Colubridae) from Yunnan Province, China, with discussion of the occurrence of D. cyanochloris (Wall, 1921) in China. Zootaxa, 4743 (1), 1 - 20. https: // doi. org / 10.11646 / zootaxa. 4743.1.1","Das, I. (2015) A Field Guide to the Reptiles of Southeast Asia. Bloomsbury Publishing, London, 376 pp.","Vogel, G. & Rooijen, J. V. (2007) A new species of Dendrelaphis (Serpentes: Colubridae) from Southeast Asia. Zootaxa, 1394 (1), 25 - 45. https: // doi. org / 10.11646 / zootaxa. 1394.1.2","Ziegler, T. & Vogel, G. (1999) On the knowledge and specific status of Dendrelaphis ngansonensis (Bourret, 1935) (Reptilia: Serpentes: Colubridae). Russian Journal of Herpelotogy, 6, 199 - 208.","Vogel, G., Rooijen, J. V. & Hauser, S. (2012) A new species of Dendrelaphis Boulenger, 1890 (Squamata: Colubridae) from Thailand and Myanmar. Zootaxa, 3392 (1), 35 - 46. https: // doi. org / 10.11646 / zootaxa. 3392.1.2","Rooijen, J. & Vogel, G. (2010) On the discovery and origin of a Javan population of the Indochinese colubrid snake Dendrelaphis subocularis (Boulenger, 1888): a multivariate study. Contributions to Zoology, 79, 85 - 92. https: // doi. org / 10.1163 / 18759866 - 07903001","Vogel, G. & Rooijen, J. V. (2011) Description of a new species of the genus Dendrelaphis Boulenger, 1890 from Myanmar (Squamata: Serpentes: Colubridae). Bonn Zoological Bulletin, 60, 17 - 24."]}

Oligodon tuani sp. nov. (Figs. 2–4) Holotype. ITBCZ 7899, subadult male, collected near Cam Ly Stream, Da Lat City, Lam Dong Province, Vietnam (coordinates 11°56’54”N, 108°24’60”E; elevation 1490 m a.s.l.) by Tuan Minh Nguyen on 15 September 2020 (Figs. 2 & 3). Paratypes. ITBCZ 7944, adult male, collected on 15 October 2020 by Tuan Minh Nguyen, coordinates 11°57’36”N, 108°23’30”E, elevation 1431 m a.s.l., from Cam Ly Road, vicinity of Da Lat City, Lam Dong Province, Vietnam; ITBCZ 7868, subadult female, collected on 12 September 2020 by Tuan Minh Nguyen, coordinates 12°01’02”N, 108°29’28”E; elevation 1520 m a.s.l., from Da Sar, Lac Duong District, vicinity of Da Lat City, Lam Dong Province, Vietnam; and ITBCZ 8215, adult female, collected from Ward no. 10, Dalat City, Lam Dong Province, Vietnam, coordinates 11°55’56”N, 108°26’58”E, elevation 1440 m a.s.l., on 24 March 2022 by Tuan Minh Nguyen (Fig. 4). Diagnosis. Oligodon tuani sp. nov. can be distinguished from its congeners by a combination of the following morphological characters: (1) large size in adults (TL ≤ 888 mm); (2) 19 dorsal scale rows at neck and midbody and 15 rows before vent; (3) TaL/TL ratio 0.184 in adult male and 0.123 in adult female; (4) ventrals 173–179 in males and 187–193 in females; (5) subcaudals 58 or 59 in males and 44 or 45 in females; (6) presubocular present; (7) 8 supralabials, fourth and fifth entering orbit; (8) cloacal plate undivided; (9) 10 maxillary teeth, posterior three enlarged; (10) vertebral stripe present; (11) temporal streak present but faint and interrupted; (12) long and deeply forked hemipenes, extending to 25 th subcaudal, without spines and papillae but with small and transversal folds and distal calyces; and (13) 13–15+3–4 dorsal blotches. Description of holotype. Subadult male, total length 585 mm; head ovoid, faintly distinct from the poorly defined neck; body robust and rather elongate; SVL 475 mm; tail quite long (TaL/TL = 0.188), 110 mm in length; eye small with round pupil, eye diameter about half of snout length. Head scalation. Rostral thick, triangular and curved on to upper snout surface, visible from above, pointed posteriorly, touching internasals, nasals and first supralabial on both sides; the portion of rostral visible from above shorter than its distance from frontal; supralabials 8/8, the 4 th and 5 th bordering eye; 1/1 loreal; 1/1 presubocular; 1/1 preocular; 2/2 postoculars; nasal divided vertically; 2+2/2+2 temporals, anterior ones elongated; prefrontal wider than long and larger than internasal; frontal large and hexagonal, slightly longer than wide (5.7 mm vs. 4.5 mm); parietals larger than the frontal, bordered laterally by the upper temporals and anteriorly by frontal, supraoculars and postoculars; 9/9 infralabials, first pair medially in contact with each other, first four in contact with anterior chin shield, the 5 th largest and touching posterior chin shield; two pairs of gular scales between posterior chin shields and preventral scale. Body scalation. Dorsal scales smooth, in 19–19–15 rows; scale row reductions from 19 to 17 at ventrals 97 and 98 and from 17 to 15 at ventrals 124 and 132; vertebral scales similar to other dorsal scales in size and shape; ventrals 173 (plus 1 preventral), angulate; cloacal plate entire; subcaudals 59, all paired; terminal caudal scale forming a pointed cap. Dentition. Maxillary teeth 10, curved posteriorly, smaller and shorter anteriorly; posterior three being enlarged and blade-shaped. Hemipenis. Hemipenis long and thin with two lobes, deeply forked at the level of 6 th SC and tip of each organ reaching the SC 25, without spines and papillae; each lobe with numerous smooth, small and transversal folds, with a deep diagonal groove; smooth hexagonal calyces on the distal part of the organ. The sulcus is prominent and divided at the level of the fork (Fig. 3). Coloration. In life, overall dorsal coloration brown to olive with 13 darker blotches on body and 4 on tail. A dark brown band from eye to eye through preoculars, supraoculars, frontal, prefrontals and internasals, continues over the 4 th and 5 th supralabials ending at lower part of the 6 th supralabial. Head with another dark brown arrowshaped marking, its apex pointing forward and reaching the posterior part of frontal, backwards obliquely crossing neck and reaching the 6 th ventral. Temporal streak faint and discontinuous, distinct posteriorly, reaching the level of ventrals. Numerous scattered dorsal black spots confined to edges of the body scales. Edge of each body scale and intervening skin yellowish to white. Vertebral stripe distinct, broader and more distinct on the anterior part of body and on the tail. Ventral color cream anteriorly to pinkish centrally and posteriorly with black rectangular blotches. Ventral surface of tail white to pinkish with black spots. In alcohol, the color became paler, but the pattern remains; venter cream to white with black blotches and dots, pinkish color disappears. Variation. Paratype ITBCZ 7944 has two preoculars on each side and its presubocular is absent on the left side. Paratype ITBCZ 8215 has 15 blotches on body and faint a vertebral stripe. Table 1 summarizes variation in size and scalation of the holotype and three paratypes. Natural history. All specimens were collected during the day. ITBCZ 7868 and 7944 were found dead on the road and the surrounding habitat is pine forest as well as flower and vegetable plantations. ITBCZ 7899 was found near Cam Ly Stream on the pine hill bordered by a vegetable plantation and residential area. ITBCZ 8215 was collected on a pine-covered hill and near a residential area. (Fig. 4E) Sexual dimorphism. Tail of adult male longer than that of adult female (TaL/TL = 0.184 vs. 0.123). Males have more subcaudals (58 or 59 vs. 44 or 45) but fewer ventrals (173–179 vs. 187–193) than females. Distribution. The new species is currently known only from Da Lat City and its vicinity, Lam Dong Province, Vietnam (Fig. 1). Etymology. We name the new species in honor of Mr. Tuan Minh Nguyen (Lam Dong Province, Vietnam) in recognition of his significant contribution to us including the type series of the new species and many other important samples of reptiles in recent years. We recommend “Langbian kukri snake” and “Rắn khiếm Langbian” for as the common English and Vietnamese names of the new species, respectively. Comparisons. Oligodon tuani sp. nov. has deeply forked hemipenis without both papillae and spines and thus differs from the following species that have conspicuous papillae on the hemipenes: O. barroni (Smith), O. deuvei David, Vogel & Rooijen, O. mouhoti (Boulenger), O. pseudotaeniatus David, Vogel & Rooijen, and O. taeniatus (Günther) (David et al. 2008b). The new species differs from O. catenatus (Blyth), O. cruentatus (Günther), O. dorsalis (Gray & Hardwicke), O. eberhardti Pellegrin, O. hamptoni Boulenger, O. planiceps (Boulenger), and O. theobaldi (Günther) by having smooth and forked (vs. spinose and unforked) hemipenis (Smith 1943). Oligodon tuani sp. nov. has 19 dorsal scale rows at midbody and thus differs from the following species that have 13 or 15 dorsal scale rows: O. annamensis Leviton, O. jintakunei Pauwels, Wallach, David & Chanhome, O. inornatus (Boulenger), O. kampucheaensis Neang, Grismer & Dattry, O. lacroixi Angel & Bourret, O. macdougalli Wall, O. nagao David, Nguyen, Nguyen, Jiang, Chen, Teynie & Ziegler, O. rostralis Nguyen, Tran, Nguyen, Neang, Yushchenki & Poyarkov, O. teyniei David, Hauser & Vogel, O. torquatus (Boulenger), and O. vertebralis Günther (Smith 1943; Vassilieva 2015; Nguyen et al. 2020; David et al. 2012, 2022). Oligodon tuani sp. nov. differs from O. albocinctus (Cantor) by having more supralabials (8 vs. 7), supralabials 4 and 5 (vs. 3 and 4) touching the eye, dorsum with blotches (vs. cross-bars), and hemipenis without papillae (with short pointed papillae) (Smith 1943); O. arenarius Vassilieva by having a larger size (TL 888 mm vs. 389 mm), more midbody scale rows (19 vs. 17), more maxillary teeth (10 vs 6–8), and more ventrals (173–193 vs. 131–144) (Vassilieva 2015); from O. booliati Leong & Grismer by having more supralabials (8 vs. 6 or 7), more ventrals (173–193 vs. 143–153), and dorsal color pattern (blotches vs. cross-bars) (Leong & Grismer 2004); from O. cattienensis Vassilieva, Geissler, Galoyan, Poyarkov, Devenger & Bohme by having more midbody scale rows (19 vs. 17), more subcaudals in males (58 or 59 vs. 32–36) and in females (44 or 45 vs. 32); fewer maxillary teeth (10 vs. 11 or 12), fewer number of dorsal blotches (13–15+3–4 vs. 24–33+5), and remarkably longer hemipenis (extending to SC 25 vs. 11 or 12) (Vassilieva et al. 2013); from O. cinereus (Günther) by having more midbody scale rows (19 vs. 17), more ventrals in females (187–193 vs. 165–185), more subcaudals (44–59 vs. 29–43), fewer maxillary teeth (10 vs. 11–14), remarkably longer hemipenis (extending to SC 25 vs. 11–14), and different dorsal color pattern (blotches vs. uniform or cross-bars or reticulations) (Smith 1943; David et al. 2012); from O. condaoensis Nguyen, Nguyen, Le & Murphy by having more midbody scale rows (19 vs. 17), more subcaudals in males (58 or 59 vs. 37) and in females (44 or 45 vs. 33–34), fewer maxillary teeth (10 vs. 11–13), remarkably longer hemipenis (extending to SC 25 vs. 13 or 14), and different dorsal color pattern (blotches vs. uniform or faint stripe) (Nguyen et al. 2016); from O. huahin Pauwels, Larsen, Suthanthangjai, David & Sumontha by having more midbody scale rows (19 vs. 17 or 15), more subcaudals in males (58 or 59 vs. 35–41), more maxillary teeth (10 vs. 6), remarkably longer hemipenis (extending to SC 25 vs. 14), and different dorsal color pattern (blotches vs. uniform or faint stripe) (Pauwels et al. 2017); from O. joynsoni (Smith) by having more midbody scale rows (19 vs. 17), fewer maxillary teeth (10 vs. 11 or 12), remarkably longer hemipenis (extending to SC 25 vs. 14), and different dorsal color pattern (blotches vs. cross-bars or reticulations) (Smith 1943; David et al. 2012, 2022); from O. macrurus (Angel) by having a shorter tail (TaL/TL = 0.184 –0.188 vs. 0.240), more midbody scale rows (19 vs. 17), more ventrals (173–193 vs. 143–152), fewer subcaudals (44–59 vs. 76–83), a shorter hemipenis (LHSC = 25 vs. 29) (Smith 1943; Geissler et al. 2011); from O. moricei David, Vogel & Rooijen by having more midbody scale rows (19 vs. 17), more ventrals (187–193 vs. 175 in females), more subcaudals (44 or 45 vs 41 in females), and dorsal blotches (vs. stripe) (David et al. 2008b); from O. phangan Pauwels, Thongyai, Chantong & Sumontha by having fewer maxillary teeth (10 vs. 12), more midbody scale rows (19 vs. 17), more ventrals (173–193 vs. 163–166), and more subcaudals (44–59 vs. 33–42) (Pauwels et al. 2021); from O. promsombuti Pauwels, Thongyai, Chantong & Sumontha by having fewer maxillary teeth (10 vs. 12), more midbody scale rows (19 vs. 17), more subcaudals (44–59 vs. 40), and remarkably longer hemipenis (extending to SC 25 vs. 13) (Pauwels et al. 2021); from O. purpurascens (Schlegel) by having faint and interrupted temporal streak (vs. distinct and continuous temporal streak), different dorsal color pattern (blotches vs. conspicuous red and banded coloration or brownish dull coloration), and forked hemipenis without papillae (vs. unforked with papillae) (Boulenger 1894; Rooijen et al. 2011; Pauwels et al. 2017; David et al. 2022); from O. saiyok Sumontha, Kunya, Dangsri & Pauwels by having fewer maxillary teeth (10 vs. 13), more midbody scale rows (19 vs. 17), more subcaudals (58 or 59 vs. 43 in males, 44 or 45 vs. 38 in females), and remarkably longer hemipenis (extending to SC 25 vs. 18) (Sumontha et al. 2017); from O. signatus (Gunther) by having more midbody scale rows (19 vs. 15–17), more ventrals (173–179 vs. 141–149 in males, 187–193 vs. 151–157 in females), and dorsal blotches (vs. cross bands or transverse rhomboidal spots) (Boulenger 1894; David et al. 2022); and from O. splendidus (Günther) by having fewer midbody scale rows (19 vs. 21), a distinct vertebral stripe (vs. vertebral stripe absent), and remarkably longer hemipenis (extending to SC 25 vs. 19) (Smith 1943). Oligodon tuani sp. nov. is similar to its congeners in the O. cyclurus species complex. However, the new species can be distinguished from O. chinensis by having more midbody scale rows (19 vs. 17), fewer subcaudals in females (44 or 45 vs. 47–53), remarkably longer hemipenis (extending to SC 25 vs. 12–14), and dorsal blotches (vs. cross bars) (David et al. 2008a; Nguyen et al. 2017); from O. culaochamensis by having a shorter tail (TaL/TL 0.184 vs. 0.218 –0.219 in males, 0.123 vs. 0.166 –0.169 in females), more ventrals in both males (173–179 vs. 167–169) and females (187–193 vs. 179–182), fewer subcaudals in both males (58 or 59 vs. 63–66) and females (44 or 45 vs. 51 or 52 in females), and remarkably longer hemipenis (extending to SC 25 vs. 20) (Nguyen et al. 2017); from O. cyclurus in having a longer tail (TaL/SVL 0.225 vs. 0.127 –0.156 in males, 0.140 vs. 0.107 –0.138 in females), more ventrals in both males (173–179 vs. 160–173) and females (187–193 vs. 168–172), more subcaudals in both males (58 or 59 vs. 37–48) and females (44 or 45 vs. 30–44 in females), and remarkably longer hemipenis (extending to SC 25 vs. 15–18) (David et al. 2008a); from O. formosanus by having a longer tail in males (TaL/SVL 0.225 vs. 0.165 –0.195) and shorter tail in females (TaL/SVL 0.140 vs. 0.149 –0.164), more subcaudals in males (58 or 59 vs. 49–55), remarkably longer hemipenis (extending to SC 25 vs. 15–18), and dorsal blotches (vs. reticulate pattern) (David et al. 2008a); from O. ocellatus by having a longer tail (TaL/SVL 0.225 vs. 0.112 –0.141 in males, 0.140 vs. 0.094 –0.114 in females), more ventrals in both males (173–179 vs. 156–165) and females (187–193 vs. 152–180), more subcaudals in both males (58 or 59 vs. 32–44) and females (44 or 45 vs. 26–33 in females), and remarkably longer hemipenis (extending to SC 25 vs. 15–17) (David et al. 2008a); and from O. saintgironsi by having a larger size (TL max 888 mm vs. 676 mm), a longer tail in males (TaL/SVL 0.225 vs. 0.191 –0.203) and shorter tail in females (TaL/SVL 0.140 vs. 0.161), more midbody scale rows (19 vs. 17 or 18), first scale reduction occurring at the level of ventrals 97–111 (vs. 62–86), more ventrals in both males (173–179 vs. 166–170) and females (187–193 vs. 184), fewer subcaudals in females (44 or 45 vs. 53), and slightly shorter hemipenis (extending to SC 25 vs. 27 or 28) (David et al. 2008a). Oligodon tuani sp. nov. is also morphologically similar to O. fasciolatus. However, it differs from O. fasciolatus by having fewer dorsal scale rows at neck (19 vs. 21–23), at midbody (19 vs. 21) and before the vent (15 vs. 17), remarkably longer hemipenis (extending to SC 25 vs. 14–21), temporal streak faint and discontinuous (vs. distinct and continuous), ventral surface with black rectangular blotches (vs. pure white or immaculate) (Gunther, 1864; David et al. 2008a; this study). The main morphological characters of the O. cyclurus species complex are summarized in Table 2.

Chai, Jing, Lu, Chen-Qi, Yi, Mu-Rong, Dai, Nian-Hua, Weng, Xiao-Dong, Di, Ming-Xiao, Peng, Yong, Tang, Yong, Shan, Qing-Hua, Wang, Kai, Liu, Huan-Zhang, Zhao, Hai-Peng, Jin, Jie-Qiong, Cao, Ru-Jun, Lu, Ping, Luo, Lai-Chun, Murphy, Robert W, Zhang, Ya-Ping, Che, Jing (2022): Discovery of a wild, genetically pure Chinese giant salamander creates new conservation opportunities. Zoological research 43 (3): 469-480, DOI: 10.24272/j.issn.2095-8137.2022.101, URL: http://dx.doi.org/10.24272/j.issn.2095-8137.2022.101

Andrias jiangxiensis sp. nov. Lu, Wang, Chai, Yi, Peng, Murphy, Zhang, and Che (Figure 4; Supplementary Table S4) Holotype: KIZ 037731, adult male (Figure 4), ex situ from first generation of offspring of the wild-caught parents, which were collected in Guanzhuang Township, Jing’an County, Jiangxi, China. Paratypes: KIZ 037728, 037730 (adult females), KIZ 037729 (adult male), all with the same collection information as the holotype. Diagnosis: Andrias jiangxiensis sp. nov. can be distinguished from its congeners by a combination of the following characters: (1) head length almost equal to width; (2) head and lower jaw relatively smooth, with small tubercles arranged irregularly; (3) lateral neck fold discontinuous with body fold at forelimb insertion; (4) finger III distinctly longer than finger I; and (5) dorsum red-brown or yellow-brown in life, with large, irregular black patches. Comparisons: Andrias jiangxiensis sp. nov. can be distinguished from A. japonicus (Temminck, 1836) by having smoother head with indistinct tubercles (vs. dense, distinct, large tubercles). Considering that no population has been genetically identified as both pure A. davidianus and A. sligoi, and that farm-bred individuals exhibit genetic contamination (Yan et al., 2018), to avoid confusion due to translocated animals or hybrids, morphological comparisons are constrained to holotypes only. Specifically, Andrias jiangxiensis sp. nov. differs from A. davidianus (holotype MNHN-RA-0.7613, images posted on: http://coldb.mnhn.fr/catalognumber/mnhn/ra/0.7613) by having a smooth head and lower jaw with indistinct, small, irregularly arranged tubercles (vs. head and lower jaw rough, with distinct, regularly arranged tubercles); from A. sligoi (holotype BMNH 1945.11.7.1., images in Turvey et al. (2019)) by having a smooth skin on the snout and around eyes (vs. with numerous distinct tubercles), and a distinct finger length formula (finger III distinctly longer than finger I vs. finger III equal to finger I). Description of holotype: Adult male, body size relatively small, SVL 381 mm, TTL 589 mm; tail muscular, thick at base, gradually flattening posteriorly and distally; dorsal fin well developed, ventral fin much reduced, maximum tail height 76.2 mm; tail short, TAL 208 mm, TAL 54.6% SVL, 35.3% TTL. Head dorsally compressed, wide, HL 102% HW; snout truncated, blunt, widen gradually posteriorly, projecting beyond lower jaw in ventral view; jaw muscular, distinctively widen from snout; jaw muscles continuous with two ovoid, convex temporal muscles dorsally posterior to each eye; nostrils small, NL 1.5 mm, anteriorly and laterally positioned, eye–nare distance 27.6 mm. Eyes small, rounded, EL 4.2 mm, 2.8 times NL, protuberant in life, lacking eyelids; dorsolaterally positioned, wide apart, IND 31.6% IOD, EL 44.2% SOD. Mouth large, corner almost reaching mid temporal protuberances; supralabial fold distinct; extending from posterior edge of mouth to axillary. Skin of head and lower jaw mostly smooth with small tubercles scattered dorsolaterally on temporal head and neck in life, all distributed irregularly, gradually increasing in numbers posteriorly; tubercles indistinct after preservation. Transverse gular fold present, without distinct glands. Body robust, dorsally compressed but less than head, elongated, AXD 34.1% TTL; costal grooves in life, less distinct after preservation; vertebral groove distinct on posterior 3/4 of body; neck fold strongly developed, from corner of mouth to posterosuperior axillary; lateral body folds strongly developed, anterior end inferior and discontinuous from neck fold at limb insertion point, extending to caudal base; single dorsolateral series of large tubercles just above base of lateral fold on each side of vertebrate from neck to caudal base, more distinct in life, barely visible after preservation; remaining body skin mostly smooth in life. Limbs short, robust, dorsoventrally compressed, with distinct lateral skin folds, more prominent on posterior limbs; hindlimbs relatively longer and stronger than forelimbs, FLL 10% TTL, PLL 12% TTL. Hands and feet dorsally compressed, finger and toe tips enlarged, rounded; finger and toe tips and ventral hands and feet covered with dark carotin layer; four fingers, no webbing, length formula II>III>I>IV, lateral skin fringe distinct on finger IV, continuous with lateral limb fold; five toes, with rudimentary webbing, toe length formula III>IV>II>V>I, toe III, IV, and Vwith stronger lateral fringes. Coloration of holotype: In life (Figure 5), dorsum red-brown, with large continuous or discontinuous irregular black patches, gradually decreasing and broken from midbody to head and tail, a few black spots scattered between large patches; ventral surface grayish brown, with black spots scattered along the mandibular margin only; small white spots with tan edge throughout the body; grayish white around eyes, iris gold with black markings. In preservative (Figure 4), bright tints faded, dorsum and abdomen faded to grayish brown and gray, respectively, white spots with tan edge, black patches and spots remain distinct; horny epidermis on tips of fingers and toes as well as palms and soles brown. Variation: No significant differences in morphology and coloration were found among all adult specimens of the type series, or between genders. However, a distinctive ontogenetic shift existed in coloration. Juveniles less than 30 cm long were mostly spotted (Figure 6A) instead of having larger patches; after exceeding 40 cm, their coloration pattern resembled the type specimens (Figure 6B). Etymology: The specific epithet “ jiangxiensis ” refers to the type locality of the new species in Jiangxi, China. It denotes the endemicity of the new species to Jiangxi based on our detailed population surveys. We suggest Jiangxi Giant Salamander as its English common name, and Ữ & 大 ª (Pinyin: Jiāng Xī Dà Ní) as its Chinese common name. Distribution and natural history: Questionnaires with local residents suggest that this species was densely distributed in two towns of Jing’an County before the 1990s. Due to commercial breeding, some indigenous purebred individuals served as breed stock in some local small farms. According to our field monitoring, the wild population of this species appears to occur only in Jiulingshan National Nature Reserve. This protected area contains intact habitat, including forested mountains and streams without villages and human activities. It also contains a diversity of teleosts, amphibians, non-avian reptiles, birds and a few mammals. During the field surveys from September 2020 to March 2022, over 291 wild individuals of adults and juveniles were found in medium-sized, pristine mountain streams with 8–10 m wide, water depth 0.1–1.5 m, and with rocky substrate (Figure 7A, B). Many natural caves formed by fallen wood, tree roots and cracks among rocks on both sides of streams provide hiding places for the new species. As a nocturnal species, A. jiangxiensis sp. nov. forages at night. Dorsal color patterning makes it blend well with the rocks at the bottom of the stream, making it difficult to find in flowing water. Vomit and excrement analyses of five wild individuals in July 2021 revealed fish, frogs, aquatic insects, and crustaceans in the natural diet. Two breeding caves were found in a backwater bay of the stream in January 2021 (Figure 7C, D) and January 2022. A large number of larvae gathered at the entrance, emerging most likely two months after hatching (Figure 7C, D). Larvae sightings peaked in February and exceeded 100 individuals per day. This continued until March, when no more new larvae emerged from the cave, and the larvae outside began to scatter throughout the stream. Combined with a questionnaire to local giant salamander farmers, we speculate that the breeding season of A. jiangxiensis sp. nov. occurs in mid-September. Conservation: Our long-term field monitoring of A. jiangxiensis sp. nov. yielded only a single incident of a breeding, wild population in the protected area at Daqi Mountain, Jing’an County. This protected area has a riverine habitat of only 6.50 km in linear distance and 36.0 km 2 of area. The upstream water runs inconstantly throughout the year, as flash floods occur occasionally in summer and droughts often occur in the winter, which lead to frequent habitat fragmentations. The downstream habitat out of the nature reserve is blocked by a dam and is inhabitable to A. jiangxiensis sp. nov. due to anthropogenic activities. This small, isolated, and sometimes fragmented population is particularly vulnerable to environmental changes, and thus the species faces a great extinction threat. Therefore, following the IUCN criteria B1 (extent of occurrence less than 100 km 2, known to exist at only a single location, and extreme fluctuation of extent of occurrence) (IUCN, 2012), we recommend to classify A. jiangxiensis sp. nov. as Critically Endangered (CR), and we recommend to list the species as Class II protected in China, and to implement further assessments. Local cultural values: Local cultural pride might have contributed positively to the conservation of indigenous species. Interviews near the survey sites found that most people did not often eat CGS, although nine out of 50 persons had eaten CGS one or two times from curiosity. Six of the 50 interviewed people in Jing’an County, including the towns Guanzhuang and Zhongyuan, could distinguish between the indigenous and introduced CGS species by the shape of head and coloration of adults. They proudly said the indigenous species is “more beautiful” than others (Supplementary Dataset S1); one person described the indigenous species as having a smoother head without tubercles. This differed from negative cultural views of CGS reported in Shaanxi (Cunningham et al., 2016), indicating diverse cultural values across China. We interviewed the owners of two small-scale farms adjacent to the field survey sites, which have existed for years (Supplementary Dataset S1). They did not cease activities following the dramatic slide in CGS prices from 3 000 RMB/kg (US $470/kg) (Cheng & Zhang, 2015; Shu et al., 2021) to 60 RMB/kg ($9/kg) after 2015. They had refused to use outside sources of CGS (Supplementary Dataset S1), even during the “salamander rush” involving large-scale salamander trading. Such small-scale farms in remote montane villages may be more likely to preserve purebred indigenous lineages than large commercial farms that frequently introduce and sell animals of unknown provenance. They can serve as effective targets for genetic screening, and as reservoirs for potential future population augmentation programs which should such be deemed necessary., Published as part of Chai, Jing, Lu, Chen-Qi, Yi, Mu-Rong, Dai, Nian-Hua, Weng, Xiao-Dong, Di, Ming-Xiao, Peng, Yong, Tang, Yong, Shan, Qing-Hua, Wang, Kai, Liu, Huan-Zhang, Zhao, Hai-Peng, Jin, Jie-Qiong, Cao, Ru-Jun, Lu, Ping, Luo, Lai-Chun, Murphy, Robert W, Zhang, Ya-Ping & Che, Jing, 2022, Discovery of a wild, genetically pure Chinese giant salamander creates new conservation opportunities., pp. 469-480 in Zoological research 43 (3) on pages 474-477, DOI: 10.24272/j.issn.2095-8137.2022.101, http://zenodo.org/record/6595559, {"references":["Yan F, Lu JC, Zhang BL, Yuan ZY, Zhao HP, Huang S, et al. 2018. The Chinese giant salamander exemplifies the hidden extinction of cryptic species. Current Biology, 28 (10): R 590 - R 592.","Turvey ST, Marr MM, Barnes I, Brace S, Tapley B, Murphy RW, et al. 2019. Historical museum collections clarify the evolutionary history of cryptic species radiation in the world's largest amphibians. Ecology and Evolution, 9 (18): 10070 - 10084.","IUCN. 2012. IUCN Red List Categories and Criteria: Version 3.1. 2 nd ed. Gland, Switzerland and Cambridge, UK: IUCN, 32.","Cunningham AA, Turvey ST, Zhou F, Meredith HMR, Guan W, Liu XL, et al. 2016. Development of the Chinese giant salamander Andrias davidianus farming industry in Shaanxi Province, China: conservation threats and opportunities. Oryx, 50 (2): 265 - 273.","Cheng YL, Zhang ZW. 2015. Analysis of the current situation of Chinese giant salamander industry in Shaanxi province. Scientific Fish Farming, 31 (9): 3 - 4. (in Chinese)","Shu GC, Liu P, Zhao T, Li C, Hou YM, Zhao CL, et al. 2021. Disordered translocation is hastening local extinction of the Chinese giant salamander. Asian Herpetological Research, 12 (3): 271 - 279."]}

During a small mammal survey in Wanglang National Natural Reserve, Sichuan, China in August–October 2018, we collected four specimens of an Asiatic striped squirrel (genus Tamiops) that could not be assigned morphologically to any described species. Phylogenetic analyses of CYTB and nuDNA sequences strongly supported them in the genus Tamiops. The p-distances of CYTB between these individuals and other species of Tamiops ranged from 17.4 to 19.1%. Morphologically, they differed from their congeners, including T. maritimus, T. mcclellandii, T. rodolphii, and T. swinhoei by pelage, skull, and baculum characteristics. Based on the results of morphological comparisons and molecular analyses, we describe the four specimens as the new species Tamiops minshanica sp. nov.

Ahaetulla rufusoculara sp. nov. Holotype. ITBCZ 7504, adult male, collected from My Phuoc Forest, My Tu District, Soc Trang Province, Vietnam; coordinates 9.56670 oN, 105.74772 oE; elevation 2 m a.s.l. by S.N. Nguyen and N.Q. Lam, on 31 December 2019 (Figs. 2 & 3). Paratypes. Five specimens: ITBCZ 7502, ITBCZ 7503, ITBCZ 7505 (subadult females) and ITBCZ 7506 (subadult male), also collected from My Phuoc Forest by the same collectors, on the same date and site as the holotype; ITBCZ 6799 (adult male) collected on 25 July 2018 by N.Q. Lam at the same site as the holotype (Figs. 4 & 5). Diagnosis. Ahaetulla rufusoculara sp. nov. is distinguished from all of its congeners by a combination of the following morphological characters: bright red eyes; medium size in adult (largest TL 1085 mm in male); snout without dermal appendage, more than two times as long as the eye diameter in adult (SnL/ED 2.33–2.37); internasal separated from supralabial by loreal; body scale rows 15-15-13; tail long (TaL/TL ratio 0.34–0.35 in males and 0.31–0.32 in females); ventrals 186–190 in males and 182–185 in females; subcaudals 143–153 in males and 128 or 129 in females; 8 (rarely 9) supralabials, fourth and fifth or fourth to sixth entering orbit; 12–14 maxillary teeth; hemipenis short, reaching 6 th or 7 th SC; cloacal plate divided; dorsum bright green; and yellow or white stripe along lower flank. Description of holotype. Adult male; head elongate and triangular (HL/HW = 2.68), distinct from neck; body elongate and compressed; SVL 682 mm; tail long (TaL/TL = 0.35), 370 mm in length; eye small, pupil horizontal, eye diameter much shorter than snout length (SnL/ED: 2.33); snout pointed without dermal appendage. Head scalation. Dorsal head scales smooth, abutting along midline rather than overlapping (Fig. 2); rostral directed forward and upward, well visible from above, touching nasals, internasals and 1 st supralabial on both sides; supralabials 8/8, 4 th, and 5 th bordering eye; 3/3 loreals; 1/1 presubocular; 1/1 enlarged preocular; 2/2 postoculars; nostril in posterior part of an elongated nasal; 2+2/2+2 temporals, with a very small scale on left side bordered by anterior temporals and postoculars; internasals longer than wide (3.0 mm in length and 1.9 mm in width), separated posteriorly from supralabials by two loreals on both sides; prefrontal bending down sharply at canthus, strongly convex, longer than wide (4.3 mm long and 2.7 mm wide), larger than internasal; frontal longest (7.6 mm long and 4.4 mm wide), widened anteriorly and much narrowed posteriorly; parietals shorter but slightly wider than frontal (6.9 mm long, 4.5 mm wide), bordered laterally by upper temporals and postocular and anteriorly by frontal and supraoculars; 9/9 infralabials, first pair medially in contact with each other, first four in contact with anterior chin shield, 5th largest and touching posterior chin shield; posterior chin shields longer than anterior ones; four pairs of gular scales between posterior chin shield and 1st ventral. Body scalation. Dorsal scales smooth, in 15-15-13 rows, lateral scales disposed obliquely, vertebral scale row slightly enlarged; scale row reduction from 15 to 13 at ventrals 149 and 150 by fusing the second and third rows of body scales on both sides; scales of sacral region strongly keeled and keel usually broken into tubercles (Fig. 3C); ventrals 190; cloacal plate entire; subcaudals 153, all paired; terminal caudal scale forming a pointed cap. Dentition. Maxillary teeth 12, forming two groups separated by a diastema: anterior group with 6 teeth, gradually enlarged posteriorly, other group with 6 teeth, first four small, last two grooved and suddenly enlarged. Hemipenis. Hemipenis short and unforked, reaching 6 th SC, with three main distinct areas: distal end papillate calyces; central part spinose with some enormous spines; and proximal area of longitudinal folds with minor scattered spines. Sulcus spermaticus sunken, not forked (Fig. 3D–F). Coloration. In life, dorsal and lateral parts of body and tail purely green; upper part of head purely green, upper lips, rostral and posterior part of canthus yellow; lower part of head yellow posteriorly and white anteriorly, except for part of 1st left and 2nd right infralabials, which are yellow; iris red; about one-third of venter yellow, gradually changing to green on remaining two-thirds of venter; ventral side of tail green; pair of longitudinal yellow parallel lines on outer margin of ventrals, extending to anterior part of tail; interstitial skin black and white, forming oblique bars, black one being fewer in number and more or less forming oblique bands when skin expanded (Figs. 2 & 3). In preservation, color faded but pattern remained: iris white; two parallel lines changing to white; all yellow parts becoming white; green faded to grey or dark brown. Variation. In adult specimens, ratio SnL/ED greater than that of subadult specimens (2.33–2.37 vs. 1.92–2.18). Paratype ITBCZ 7502 with small scale divided from upper part of 2 nd and 3 rd supralabials; suture between them forms a Y-shaped slit. Paratype ITBCZ 7503 with 9 supralabials, 4 th –6 th entering eye on both sides. Left side 5 th and 6 th supralabials not separated completely from each other, their suture not reaching edge of upper lip. Paratype ITBCZ 7505 with 2 loreals on right side and no presubocular on left side; 10 th and 14 th ventrals divided partially by transverse suture half width of given ventral. Paratype ITBCZ 7506 with incompletely divided precloacal plate. Paratypes ITBCZ 7502, ITBCZ 7505, ITBCZ 7506, and ITBCZ 6799 with 2 preventrals, posterior one contacting 1st ventral divided. Table 4 summarizes variation in size and scalation of type series. Field notes. All specimens were collected at night, from 7:00 PM to 1:00 AM, in open areas along channel banks in wetland forest dominated by cajuput samet tree (Melaleuca cajuputi Powell) (Fig. 6). The snakes were sleeping on bushes at 1–2 m above ground. Congener Ahaetulla cf. nasuta was found sympatrically. Sexual dimorphism. Males (n = 3) have longer relative tails than those of females (n = 3) (TaL/TL 0.34–0.35 in males; 0.31–0.32 in females) and more subcaudals (SC 134–153 in males, 128 or 129 in females) (Table 4). Keeled scales on sacral region present on males only. In life, males have venter yellow anteriorly and green posteriorly while venter of females is white with light green blotches posteriorly. Distribution. The new species is currently known only from My Phuoc Forest, My Tu District, Soc Trang Province, Vietnam (Fig. 6). Etymology. The specific epithet rufusoculara refers to the red color of the eyes and is derived from the Latin adjective rufus (meaning “red”) and oculus (means “eye”). We recommend “Red-eyed vine snake” and “Rắn roi mắt đȯ” as the common English and Vietnamese names of the new species, respectively. Comparisons. Ahaetulla rufusoculara sp. nov. differs morphologically from all congeners as follows: from A. anomala Annandale, A. borealis Mallik, Srikanthan, Pal, D’Souza, Shanker & Ganesh, A. farnsworthi Mallik, Srikanthan, Pal, D’Souza, Shanker & Ganesh, A. isabellina (Wall), A. laudankia Deepak, Narayanan, Sarkar, Dutta & Mohapatra, A. malabarica Mallik, Srikanthan, Pal, D’Souza, Shanker & Ganesh, A. nasuta, A. oxyrhyncha (Bell), A. pulverulenta (Dumeril, Bibron & Dumeril), and A. sahyadrensis Mallik, Srikanthan, Pal, D’Souza, Shanker & Ganesh by the absence (vs. presence) of dermal appendage on snout (Deepak et al. 2019; Mallik et al. 2020); from A. dispar (Gunther) by having more ventrals (182–190 vs. 136–156) and more subcaudals (128–153 vs. 84–119) (Smith 1943); from A. fasciolata (Fischer) by having fewer ventrals (182–190 vs. 221–231), fewer subcaudals (128–153 vs. 185–194), and precloacal divided (vs. entire) (Boulenger 1896); from A. fronticincta (Gunther) by having nasals distinctly separated from each other (vs. nasals usually in contact with one another above the rostral), 3 rd and 4 th supralabials not horizontally divided (vs. horizontally divided), and 2 or 3 supralabials (vs. 1) entering eye (Boulenger 1896; Smith 1943); from A. mycterizans by having fewer ventrals in males (186–190 vs. 192–194), fewer subcaudals (143–153 vs. 167 or 168 in males, 128 or 129 vs. 132–156 in females), precloacal divided (vs. entire), and slightly longer hemipenes (extending to SC 6 or 7 vs. 5) (Linnaeus 1758; Smith 1943; Miralles & David 2010); from A. perroteti (Dumeril, Bibron & Dumeril) by having more ventrals (182–190 vs. 136–146), more subcaudals (143–153 vs. 65–75 in males and 128 or 129 vs. 71–168 in females), and presence (vs. absence) of 2 or 3 loreals (Smith 1943); from A. prasina by having fewer ventrals (182–190 vs. 194–235) and fewer subcaudals (143–153 vs. 165–187 in males and 128 or 129 vs. 151–172 in females) (Smith 1943); and from A. travancorica Mallik, Srikanthan, Pal, D’Souza, Shanker & Ganesh by having more ventrals (186–190 vs. 153 in males) and more subcaudals (143–153 vs. 130 in males) (Mallik et al. 2020)., Published as part of Lam, Ngon Quang, Thu, Tran Thi Anh, Nguyen, Luan Thanh, Murphy, Robert W. & Nguyen, Sang Ngoc, 2021, A new species of Ahaetulla Link, 1807 (Squamata: Colubridae) from the Mekong Delta, Vietnam, pp. 290-304 in Zootaxa 4966 (3) on pages 296-301, DOI: 10.11646/zootaxa.4966.3.2, http://zenodo.org/record/4736630

Gekko phuyenensis sp. nov. English: Phu Yen Marbled Gecko Vietnamese: Tắc kè phú yên Figs. 2, 3. Holotype. ITBCZ 6653, adult male (Figs. 2, 3A), Ninh Dong Commune, Tuy An District, Phu Yen Province, Vietnam, at an elevation of 5 m above sea level (13˚21’N, 109˚17’E), collected by V.D.H. Nguyen on 16 July 2018. Paratypes. ITBCZ 6654 and ITBCZ 6656 (adult males), and ITBCZ 6655 (adult female, Fig. 3B), the same data as the holotype. Diagnosis. Gekko phuyenensis sp. nov. is distinguished from all of its congeners by a combination of the following morphological characters: medium size in adults (SVL up to 92.8 mm; nares in contact with rostral; nasals 3; internasal 1; interorbitals 35 or 36; ventrals 30–32; dorsal tubercle rows 9 or 10; precloacal pores (in males) 12 or 13, precloacal pits (in female) 13; subdigital lamellae under first finger 18–21, fourth finger 19 or 20, first toe 18–20, and fourth toe 21–23; subcaudals median enlarged; no webbing between toes, and absence of tubercles on hind limbs. Description of holotype. Adult male. Size moderate, SVL 91.8 mm, tail original, TaL 104.5 mm; head relatively long (HL/SVL 0.29), depressed (HH/HL 0.40), distinct from neck; snout elongate (SE/HL 0.43), longer than eye diameter (ED/SE 0.46); scales on dorsal head and snout small, granular, and homogeneous; interorbitals 35; nares closer to snout-tip than to eye (NE/SE 0.78), bordered by rostral, nasorostral, supranasal, postnasal, and first supralabial; internasal 1; eyes medium in size (ED/HH 0.5), pupil vertical, superciliaries smooth, 8 ciliary spines; ear opening small (EarD/HL 0.15) and obliquely oval; eye to ear distance larger than eye diameter (EE/ED 1.48); supralabials 12 and infralabials 10, followed by small scales extending to jaw; rostral wider than mental; gulars 4; flat granules dominantly cover gular region. Body elongate (AG/SVL 0.48) with weak ventrolateral folds; scales on back small, round, granular, and intermixed with 10 enlarged and smooth dorsal tubercle rows at midbody; tubercles absent on forelimbs, hindlimbs, throat, and ventolateral folds; dorsal tubercles surround by 9 small granule scales; ventrals smooth and larger than dorsal scales; 32 scale rows across belly between midbody ventolateral folds; scales along underside of body from mental to front of cloacal slit 155; precloacal pores 12, arranged in two angular series interrupted by a poreless scale; 9 enlarged scale rows between pores and cloacal slit (Fig. 3A). Limbs relatively long (ForeaL/SVL 0.14 and CrusL/SVL 0.18); scales on limbs homogeneously round and granular, without tubercles; fingers and toes compressed and without webbing between digits; claw absent on inner fingers and toes (Fig. 2C–D); subdigital lamellae under first finger 21, fourth finger 20, first toe 18, and fourth toe 22. Original tail longer than snout-vent length (TaL/SVL 1.14), thicker at base; scales on dorsum of tail small, round, and granular; 99 median enlarged subcaudals; tail base with tubercles on dorsal surface and 2 enlarged smooth postcloacal tubercles on each ventrolateral side. In everted condition, hemipenis smooth and distinctly bifurcated near tip forming 2 symmetric lobes; on asulcal side, proximal part smooth while tiny calyces occur from middle of organ to lobes (Fig. 2E); on sulcal side, sulcus lips prominent and smooth while lobes bear distinct folds (Fig. 2F). Coloration. In life, dorsal background dark green to dark brown. Dorsum with darkish blotches and smaller whitish blotches alternately. Blotches enlarged on vertebrate and symmetrically but smaller on flanks. Ventral side yellowish to white without pattern. In preservative, the color became faint, but pattern is constant (Fig. 2 A&B). Variation. Paratypes agree with holotype in color pattern both in life and in preservation. Maximum SVL is 92.8 mm in paratype ITBCZ 6654 (male). Series of precloacal pores of two males, including the holotype (ITBCZ 6653) and paratype (ITBCZ 6654), are interrupted by a pore-less scale, while they are continuous in paratype (ITBCZ 5566). Variation in size and scalation of the type series are shown in Table 3. Sexual dimorphism. The adult female slightly smaller than adult males (SVL 90.2 mm versus 91.8–92.8 mm [n = 3]). The female with 13 precloacal pits (Fig. 3B) while males bear distinct pores (Fig. 3). Etymology. The specific epithet phuyenensis is a toponym derived from the Phu Yen Province, southern Vietnam where the new gecko was discovered. Comparisons. Among members of the Gekko petricolus group, Gekko phuyenensis sp. nov. can be distinguished from G. badenii by having fewer dorsal tubercles rows (9 or 10 versus 12–17), more subdigital lamellae beneath first and fourth toes (18–20 versus 12–14 and 21–23 versus 15–20, respectively), absence (versus presence) of back banded, and presence (versus absence) of precloacal pits in the female; from G. boehmei by having fewer dorsal tubercles rows (9 or 10 versus 12–15), absence (versus weakly developed) of toe webbing, absence (versus presence) of tubercles on hind limbs, presence (versus absence) of precloacal pits in the female, and higher number of subdigital lamellae under first and fourth fingers, first and fourth toes (18–21 versus 12 or13, 19 or 20 versus 15 or 16, 18–20 versus 14–16, and 21–23 versus 17 or 18, respectively); from G. canaensis by having more subdigital lamellae under first and fourth fingers, first and fourth toes (18–21 versus 14–17, 19–20 versus 16–18, 18–20 versus 14–16, and 21–23 versus 17–19, respectively) and more precloacal pits in the female (13 versus 0–3); from G. grossmanni by having more subdigital lamellae under fourth fingers, first and fourth toe (19 or 20 versus 17 or 18, 18–20 versus 15 or 16, and 21–23 versus 19 or 20, respectively) and more precloacal pits in the female (13 versus 0–2); from G. cf. grossmanni in Binh Hung, Khanh Hoa Province, Vietnam (n=1) by having fewer dorsal tubercle rows (9–10 versus 12), presence (versus absence) of internasal, and more subdigital lamellae under fourth fingers, first and fourth toes (19 or 20 versus 17, 18–20 versus 16, and 21–23 versus 16, respectively); from G. cf. grossmanni in Hon Ba, Khanh Hoa Province, Vietnam (n=3) by fewer dorsal tubercle rows (9 or 10 versus 12 or 13), presence (versus absence) of internasal, and more subdigital lamellae under first and fourth fingers, first and fourth toes (18–21 versus 17, 19 or 20 versus 17 or 18, 18–20 versus 17, and 21–23 versus 18 or 19, respectively); from G. cf. grossmanni in Ninh Hoa, Khanh Hoa Province, Vietnam (n=3) by having a larger SVL (90.2–92.8 versus 70.2–76.4 mm), fewer interorbitals (35 or 36 versus 42–44), more granule surrounding dorsal tubercle (9 versus 8), more subdigital lamellae under fourth fingers, first and fourth toes (19 or 20 versus 15, 18–20 versus 14–16, and 21–23 versus 16–18, respectively); from G. flavimaritus by having a larger SVL (90.2–92.8 versus 67.5–84.5 mm), fewer dorsal tubercles rows (9 or 10 versus 12–16), more subdigital lamellae under first and fourth toes (18–20 versus 10–15, and 21–23 versus 15–18, respectively), absence (versus weakly developed) of toe webbing, more precloacal pores in males (12 or 13 versus 7 or 8) and presence (versus absence) of precloacal pits in the female; from G. lauhachindai by having fewer dorsal tubercles rows (9 or 10 versus 14), absence (versus presence) of back banded, presence (versus absence) of precloacal pits in the female, and more subdigital lamellae under first and fourth toes (18–20 versus 13 and 21–23 versus 13–15, respectively); from G. petricolus by having fewer dorsal tubercles rows (9 or 10 versus 15–18), more subdigital lamellae under first and fourth toes (18–20 versus 12–14 and 21–23 versus 16–18, respectively), and absence (versus presence) of tubercles on hindlimbs; from G. russelltraini by having fewer dorsal tubercles rows (9 or 10 versus 12–16), more subdigital lamellae under first and fourth fingers, first and fourth toes (18–21 versus 12–16, 19 or 20 versus 14–17, 18–20 versus 15 or 16, and 21–23 versus 17–19, respectively), more precloacal pores in males (12 or 13 versus 8–11) and presence (versus absence) of precloacal pits in the female; and from G. takouensis by having fewer dorsal tubercles rows (9 or 10 versus 14–17), more subdigital lamellae under first and fourth fingers, first and fourth toes (18–21 versus 15, 19 or 20 versus 17 or 18, 18–20 versus 14–16, and 21–23 versus 18–20, respectively), and presence (versus absence) of precloacal pits in the female. Distribution. Gekko phuyenensis sp. nov. is currently known from the type locality in Ninh Dong Commune, Tuy An District, Phu Yen Province, Vietnam (Fig. 1). Natural history. All specimens were collected during the daytime, on rocky outcrop walls close to the sea (Fig. 4). Surrounding habitat was shrubs, liana, interspersed by outcrop granite.At the collected sites, the temperature was 28.1–31.4 oC, relative humidity was 62.3–67.9%.

Subdoluseps vietnamensis sp. nov. (Figs. 3 & 4) Holotype. ITBCZ 5842, adult male, collected from Ho Tram Area, Xuyen Moc District, Ba Ria-Vung Tau Province, Vietnam; coordinates 10��29���38.2��� N, 107��27���40.0��� E; elevation 11 m asl by Sang N. Nguyen and Vu D.H. Nguyen, on 21 March 2017 (Fig. 3). Paratypes. Two adult males ITBCZ 5843 and 5844, and one adult female ITBCZ 5856 collected by Sang N. Nguyen and Vu D.H. Nguyen from the same site as the holotype; two adult males, ITBCZ 7038 and 7046, collected by Manh V. Le on 04 June 2020 at Ho Tram Area, Xuyen Moc District, Vietnam, coordinates 10��28���59.8��� N, 107��26���03.6��� E, elevation 10 m asl; two adult females ITCBZ 7056 and 7059, and one juvenile ITBCZ 7070, collected by Manh V. Le on 04 June 2020 at La Gi Town, Binh Thuan Province, Vietnam, coordinates 10��42���38.8������ N, 107��42���10.8��� E, elevation 62 m asl (Fig. 4). Diagnosis. Subdoluseps vietnamensis sp. nov. is distinguished from its congeners by a combination of the following morphological characters: medium size in adults (SVL up to 48.7 mm); tail length approximately equal to SVL; 27���30 smooth midbody scale rows; dorsal scales not enlarged; 55���57 paravertebral scales; 34���37 axilla-groin scales; 55���62 ventral scale rows; 64���74 subcaudal scales; four supraoculars; frontoparietal single; prefrontals not in contact with one another; two loreal scales; seven supralabials, the fifth below the center of the eye; one anterior and two posterior enlarged temporal scales; ear opening with two lobules on the anterior margin; 9 or 10 smooth subdigital lamellae beneath finger III and 12���15 beneath toe IV; six enlarged precloacal scales; and four distinct black stripes on dorsum. Description of holotype. Adult male, SVL 46.5 mm; snout short and obtuse; lower eyelid scaly; body rather robust; tail regenerated. Head scales smooth; rostral convex, distinctly visible from above, broader than long (2.0 mm width, 1.0 mm height); a pair of supranasals forming short median suture, touching nasals and anterior loreals laterally, width equal to length (0.7 mm); prefrontals small, quadrangular, widely separated from each other, each touching both loreals; four supraoculars; frontal truncate anteriorly, longer than its distance from tip of snout, shorter than its distance from nuchal (2.0 mm vs. 3.2 mm), longer than wide (1.8 mm width, 2.6 mm length), touching two anterior supraoculars; frontoparietal single, much larger than interparietal, wider than long (3.0 mm width, 1.9 mm length), touching three supraoculars; parietals in contact posteriorly behind the interparietal; interparietal longer than wide (1.1 mm width, 1.6 mm length), with parietal eye visible posterior to center; a pair of nuchals; seven supralabials on both sides, fifth below center of the eye, first larger than the three following, sixth largest; two loreals, anterior (0.6 mm length, 0.6 mm height) slightly higher, the posterior (0.9 mm length, 0.5 mm height) longer; nasal divided, in contact with the first supralabial, rostral, anterior loreal, and supranasal; nostril in center of nasal; eight supraciliaries, first largest; one enlarged anterior temporal in contact with the sixth and the seventh supralabial; two posterior temporals, lower one smaller and overlapping the upper one, the lower temporal in contact with seventh supralabials; six infralabials, first two in contact with postmental; two pairs of chin shields, first pair medially in contact with each other; ear-opening moderately small, the anterior border with two lobules. Body elongate (AGD/SVL = 0.56); dorsal scales smooth, not larger than lateral and ventral scales, with eight longitudinal rows on the back; 28 midbody scale rows; 71 paravertebral scales; 35 axilla-groin scales; ventral scales smooth, in 55 rows; limbs short (FLL/SVL = 0.18, HLL/SVL = 0.25), pentadactyl, widely separated by ten scales when adpressed; 3, 6, 9, 9, and 7 smooth lamellae beneath fingers I���V, respectively; 4, 7, 13, 13, and 9 smooth lamellae beneath toes I���V, respectively; six enlarged precloacal scales, the two middle ones slightly larger than the others; size of median subcaudal series on the original part of the tail equal to those of its adjacent rows. Coloration. In life, overall dorsal coloration reddish brown with two dorsolateral bright lines on anterior part of dorsum; each dorsal scale of four central rows with a black spot forming four distinct longitudinal lines on dorsum; ear and surrounding area orange; sides dark brown to black intermixed with white, brown and yellow spots; chest and belly yellow (Fig. 3). In preservation, color fades but the four longitudinal dorsal lines remain distinct; orange and yellow faded to cream or white; overall dorsal and lateral coloration black to dark brown with bright spots on lateral sides; venter cream. Variation in paratypes (n = 8). Most morphological characters of paratypes agree with those of the holotype. The following main characters vary: (1) midbody scale rows: vary from 27 to 30; (2) paravertebral and ventral scale rows: range from 55 to 57 and from 55 to 62, respectively; (3) axilla-groin scales: range from 34 to 37; (4) and nuchals: range from one to three scales. Table 3 summarizes variation in size and scalation of the type series. ...Continued on the next page ...Continued on the next page Field notes. All specimens were collected during the daytime, on sandy ground among rotting leaves in Earleaf acacia (Acacia auriculiformis A.Cunn. ex Benth.) plantation, secondary dipterocarp forest, and rubber plantation in lowland coastal areas (Fig. 5). The holotype and paratypes ITBCZ 5843 & 5844 were collected at 15:42, air temperature 30.3 oC, relative humidity 70.2 %; ITCBZ 7038 was collected at 13:30, air temperature 33 oC, relative humidity 75 %; ITBCZ 7046 was collected at 8:54, air temperature 30 oC, relative humidity 80 %; ITBCZ 7056, 7059 & 7070 were collected at 12:10, air temperature 32 oC, relative humidity 65%. To avoid capture, this species can dive into and stay under loose sand. Sexual dimorphism. Adult males mainly differ from adult females in body coloration in life and shape of tail base. Males have yellow color on chest and belly (vs. white, cream or yellowish color in females), orange around the ear (vs. dark brown in females), and distinct bright spots on lateral side (vs. pure dark brown or with small bright dots in females). Ventral side of tail base is slightly swollen in males and flat in females (Figs. 3, 4A���E). Juvenile has bright golden color on the dorsum with four clear dorsal longitudinal stripes (Fig. 4F). Distribution. The new species is currently known only from (1) Ho Tram, Xuyen Moc District, Ba Ria-Vung Tau Province and (2) La Gi Town, Binh Thuan Province, southern Vietnam (Fig. 1). Etymology. The new skink is named after the nation of Vietnam. We recommend ��� Vietnam Agile Skink��� and ���Th���n l���n ch��n ng���n vi���t nam��� as the common English and Vietnamese names of the new species, respectively. Comparisons. Subdoluseps vietnamensis sp. nov. differs morphologically from other species of the Lygosoma s.l. in Asia as follows: For Subdoluseps, S. vietnamensis sp. nov. differs from S. herberti, S. malayana, S. purthi, and S. samajaya by having a single frontoparietal scale (vs. paired) (Grismer 2019; Karin et al. 2018; Sharma 1977; Smith 1916). In addition, it differs from S. herberti by having a smaller adult size (maximum SVL = 48.7 mm vs. 54.0 mm), presence (vs. absence) of nuchal scales (1���3 vs. 0), and smooth dorsal scales (vs. dorsal with 5 strong keels) (Smith 1916); from S. malayana by having a smaller adult size (maximum SVL = 48.7 mm vs. 65.4 mm), more supralabials (7 vs. 6) with the fifth below the center of the eye (vs. the fourth below the center of the eye), smooth dorsal scales (vs. dorsal with 3���5 strong keels) (Grismer et al. 2019); from S. purthi by having a smaller adult size (maximum SVL = 48.7 mm vs. 67.0 mm), fewer midbody scale rows (27���30 vs. 32���34), and more paravertebral scale rows (55���57 vs. 50) (Sharma 1977); from S. samajaya by having a smaller adult size (maximum SVL = 48.7 mm vs. 70.1 mm), fewer paravertebral scale rows (55���57 vs. 60 or 61), and smooth dorsal scales (vs. dorsal with 5 strong keels) (Karin et al. 2018). Subdoluseps vietnamensis sp. nov. is phenotypically most similar to S. bowringii and S. frontoparietale but can be distinguished from both based on a combination of characters. The new species differs from S. bowringii by having a single frontoparietal scale (vs. paired) (Geissler et al. 2011; Smith 1935; this study), tail length approximately equal to SVL (vs. tail longer than SVL) (mean TL/SVL = 1.04 vs. 1.27 [Smith 1935; this study]), and fewer subcaudal scales (64���74 vs. 76���81) (this study); from S. frontoparietale by having a tail length approximately equal to SVL (mean TL/SVL = 1.04 vs. tail longer than SVL, mean TL/SVL = 1.24 [Taylor 1962; this study, Table 4]), fewer subcaudal scales (64���74 vs. 78���80) [Taylor 1962; this study]), fewer axilla-groin scales (34���37 vs. 39���42) (this study), four distinct longitudinal black stripes on dorsum (vs. six distinct black stripes) (Taylor 1962; this study), and absence (vs. presence) of bright dorsolateral band extending from eye to tail base (Taylor 1962; this study, Fig. 6). From genus Lygosoma s.s., Subdoluseps vietnamensis sp. nov. differs from L. isodactylum, L. siamensis, L. tabonorum, and L. quadrupes (Linnaeus) by having supranasals in contact with each other (vs. not in contact) (Geissler et al. 2011, Heitz et al. 2016, Siler et al. 2018) and from L. corpulentum by having smaller adult size (maximum SVL = 48.7 mm vs. 170.0 mm), frontoparietal scale single (vs. paired), and fewer midbody scale rows (27���30 vs. 36���40) (Smith 1921; Geissler et al. 2011). For the genus Riopa, Subdoluseps vietnamensis sp. nov. differs from R. albopunctata, R. anguinua, R. goaensis, R. guentheri, R. popae Shreve, and R. punctata (Gmelin) by having frontoparietal scales single (vs. paired) (Geissler et al. 2012; Sharma 1976; Smith 1935); from R. lineata and R. lineolata by having fewer paravertebral scale rows (55���57 vs. 104���110 in R. lineata [Smith 1935] and 78���93 in R. lineolata [Siler et al. 2018]); and from R. vosmaeri by having hind-limb with 5 (vs. 4) toes (Gray 1939; Seetharamaraju et al. 2009). For remaining Asian species of the Lygosoma s.l. that were not confirmed by Freitas et al. (2019), Subdoluseps vietnamensis sp. nov. differs from L. boehmei Ziegler, Schmitz, Heidrich, Vu & Nguyen, L. opisthorhodum Werner, L. singha (Taylor), and L. veunsaiensis Geissler, Hartmann & Neang by having frontoparietal scale single (vs. paired) (Geissler et al. 2012; Werner 1910; Ziegler et al. 2007); from L. angeli (Smith) by having a smaller adult size (maximum SVL = 48.7 mm vs. 100.0 mm) and fewer paravertebral scale rows (55���57 vs. 107���115) (Smith 1937; Geissler et al. 2011); from L. haroldyoungi (Taylor) by having a smaller adult size (maximum SVL = 48.7 mm vs. 136.0 mm), fewer paravertebral scale rows (55���57 vs. 141���145), and fewer midbody scale rows (27���30 vs. 38���42) [Taylor 1962; Geissler et al. 2011]; from L. kinabatanganensis Grismer, Quah, Duzulkafly & Yambun by having a smaller adult size (maximum SVL = 48.7 mm vs. 141.0 mm), fewer paravertebral scale rows (55���57 vs. 98), and fewer midbody scale rows (27���30 vs. 42) (Grismer et al. 2018); from L. koratense Smith by having a smaller adult size (maximum SVL = 48.7 mm vs. 105.0 mm) and fewer midbody scale rows (27���30 vs. 32���34) [Smith 1935]; and from L. bampfyldei Bartlett, L. peninsulare Grismer, Quah, Duzulkafly & Yambun, and L. schneideri Werner by having fewer paravertebral scale rows (55���57 vs. 81���85 in L. bampfyldei [Grismer et al. 2018], 87 in L. peninsulare [Grismer et al. 2018], and 95 in L. schneideri [Grismer et al. 2018])., Published as part of Le, Manh Van, Nguyen, Vu Dang Hoang, Phan, Hoa Thi, Rujirawan, Attapol, Aowphol, Anchalee, Vo, Thi-Dieu-Hien, Murphy, Robert W. & Nguyen, Sang Ngoc, 2021, A new skink of the genus Subdoluseps Freitas, Datta-Roy, Karanth, Grismer & Siler, 2019 (Squamata: Scincidae) from southern Vietnam, pp. 257-274 in Zootaxa 4952 (2) on pages 263-270, DOI: 10.11646/zootaxa.4952.2.3, http://zenodo.org/record/4674021, {"references":["Taylor, E. H. (1962) New oriental reptiles. The University of Kansas Science Bulletin, 43, 209 - 263. https: // doi. org / 10.5962 / bhl. part. 13346","Grismer, L. L., Dzukafly, Z., Muin, M. A., Quah, E. S., Karin, B. R., Anuar, S. & Freitas, E. S. (2019) A new skink of the genus Subdoluseps (Hardwicke & Gray, 1828) from Peninsular Malaysia. Zootaxa, 4609 (2), 358 - 372. https: // doi. org / 10.11646 / zootaxa. 4609.2.10","Karin, B. R., Freitas, E. S., Shonleben, S., Grismer, L. L., Bauer, A. M. & Das, I. (2018) Unrealized diversity in an urban rainforest: A new species of Lygosoma (Squamata: Scincidae) from western Sarawak, Malaysia (Borneo). Zootaxa, 4370, 345 - 362. https: // doi. org / 10.11646 / zootaxa. 4370.4.2","Sharma, R. C. (1977) A new lizard of the genus Riopa Gray (Scincidae) from Tamil Nadu, India. Records Zoological Survey of India, 73, 41 - 42.","Smith, M. A. (1916) Description of three new lizards and a new snake from Siam. The journal of the Natural History Society of Siam, 2, 44 - 47.","Geissler, P., Nguyen, Q. T., Phung, M. T., Devender, R. W. T., Hartmann, T., Farkas, B., Ziegler, T. & Bohme, W. (2011) A review of Indochinese skinks of the genus Lygosoma Hardwicke & Gray, 1827 (Squamata: Scincidae), with natural history notes and an identification key. Biologia, 66, 1159 - 1176. https: // doi. org / 10.2478 / s 11756 - 011 - 0130 - 2","Smith, M. A. (1935) The fauna of British India, Including Ceylon and Burma. Reptiles and Amphibia, Vol. II. Sauri a. Taylor and Francis, London, 440 pp.","Heitz, B. B., Diesmos, A. C., Freitas, E. S., Ellsworth, E. D., Grismer, L. L., Aowphol, A., Brown, R. M. & Siler, C. D. (2016) New supple skink, genus Lygosoma (Reptilia: Squamata: Scincidae) from the Palawan Faunal Region, Philippines. Herpetologica, 72, 352 - 361. https: // doi. org / 10.1655 / Herpetologica-D- 16 - 00023.1","Siler, C. D., Heitz, B. B., Davis, D. R., Freitas, E. S., Aowphol, A., Termprayoon, K. & Grismer, L. L. (2018) New Supple Skink, Genus Lygosoma (Reptilia: Squamata: Scincidae), from Indochina and Redescription of Lygosoma quadrupes (Linnaeus, 1766). Journal of Herpetology, 52, 332 - 347. https: // doi. org / 10.1670 / 16 - 064","Smith, M. A. (1921) New or little - known Reptiles and Batrachians from Southern Annam (Indochina). Proceedings of the Zoological Society of London, 423 - 440. https: // doi. org / 10.1111 / j. 1096 - 3642.1921. tb 03271. x","Geissler, P., Hartmann, T. & Neang, T. (2012) A new species of the genus Lygosoma Hardwicke & Gray, 1827 (Squamata: Scincidae) from northeastern Cambodia, with an updated identification key to the genus Lygosoma in mainland Southeast Asia. Zootaxa, 3190 (1), 56 - 68. https: // doi. org / 10.11646 / zootaxa. 3190.1.4","Sharma, R. C. (1976) Records of the reptiles of Goa. Records Zoological Survey of India, 71 (1975), 149 - 167.","Seetharamaraju, M., Sreekar, R., Srinivasulu, C., Srinivasulu, B., Kaur, H. & Venkateshwarlu, P. (2009) Rediscovery of Vosmer's Writhing Skink Lygosoma vosmaerii (Gray, 1839) (Reptilia: Scincidae) with a note on its taxonomy. Journal of Threatened Taxa, 1, 624 - 626. https: // doi. org / 10.11609 / JoTT. o 2160.624 - 6","Freitas, E. S., Datta-Roy, A., Karanth, P., Grismer, L. L. & Siler, C. D. (2019) Multilocus phylogeny and a new classification for African, Asian and Indian supple and writhing skinks (Scincidae: Lygosominae). Zoological Journal of the Linnean Society, 186, 1067 - 1096. https: // doi. org / 10.1093 / zoolinnean / zlz 001","Werner, F. (1910) Uber neue oder seltene Reptilien des Naturhistorischen Museums in Hamburg. II. Eidechsen. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 2, 1 - 46.","Ziegler, T., Schmitz, A., Heidrich, A., Vu, N. T. & Nguyen, Q. T. (2007) A new species of Lygosoma (Squamata: Sauria: Scincidae) from the central Truong Son, Vietnam, with notes on its molecular phylogenetic position. Revue Suisse de Zoologie, 114, 397 - 415. https: // doi. org / 10.5962 / bhl. part. 80396","Smith, M. A. (1937) Un nouveau Lezard de Cochinchine. Bulletin du Museum National d'Histoire Naturelle, 9, 366.","Grismer, L. L., Quah, E. S. H., Zaharil, D. & Paul, Y. (2018) On the taxonomy of Lygosoma bampfyldei Bartlett, 1895 (Squamata: Scincidae) with descriptions of new species from Borneo and Peninsular Malaysia and the resurrection of Lygosoma schneideri Werner, 1900. Zootaxa, 4438, 528 - 550. https: // doi. org / 10.11646 / zootaxa. 4438.3.6"]}

Le, Manh Van, Nguyen, Vu Dang Hoang, Phan, Hoa Thi, Rujirawan, Attapol, Aowphol, Anchalee, Vo, Thi-Dieu-Hien, Murphy, Robert W., Nguyen, Sang Ngoc (2021): A new skink of the genus Subdoluseps Freitas, Datta-Roy, Karanth, Grismer & Siler, 2019 (Squamata: Scincidae) from southern Vietnam. Zootaxa 4952 (2): 257-274, DOI: https://doi.org/10.11646/zootaxa.4952.2.3

Scincella baraensis sp. nov. (Figs. 2, 3) Holotype. ITBCZ 6534, adult male, collected from Ba Ra Mountain, Binh Phuoc Province, Vietnam; coordinates 11°49’42.0” N, 107°00’30.8” E; elevation 337 m a.s.l. by Sang N. Nguyen and Vu D.H. Nguyen, on 4 June 2018 (Fig. 2). Paratypes. Two specimens, also collected from Ba Ra Mountain by Sang N. Nguyen and Vu D.H. Nguyen on the same date and coordinates as the holotype: ITBCZ 6535, adult male (Fig. 3 A–C) and ITBCZ 6536, adult female (Fig. 3D). Diagnosis. Scincella baraensis sp. nov. is distinguished from all of its congeners by a combination of the following morphological characters: relatively small size in adults (SVL up to 49.2 mm); when limbs adpressed toes do not or just reach fingers; 30 smooth midbody scale rows; dorsal scales not enlarged; 66–70 paravertebrals; 64–66 ventral scale rows; 4 supraoculars; prefrontals in broad contact with or separated from one another; 2 loreals; 7 supralabials, the fifth and sixth below the center of the eye; 1 anterior and 2 posterior enlarged temporals; 3 or 3.5 pairs of nuchals; tympanum deeply sunk with weak auricular lobules on anterior margin of ear; 12 or 13 smooth lamellae beneath finger IV and 18–20 beneath toe IV; 2 enlarged precloacals; hemipenis smooth, short, unforked; dorsum with faint black dots. Description of holotype. Adult male; SVL 46.8 mm; snout short and obtuse; lower eyelid with an undivided transparent disc; body rather compressed; tail longer than snout-vent length, regenerated; limbs pentadactyl, toes just reach to fingers when limbs adpressed. Head scales smooth; rostral convex, distinctly visible from above, in broad contact with frontonasal, broader than long (1.8 mm width, 0.8 mm height); no supranasals; prefrontals in broad contact with one another (1.1 mm width, 1.1 mm length, in contact 0.4 mm); four supraoculars; a pair of frontoparietals, shorter than frontal; frontal narrowing posteriorly, longer than wide, bordered laterally by first two supraoculars, anteriorly by prefrontals, and posteriorly by frontoparietals; frontoparietals in contact with supraoculars 2–4; parietals in contact posteriorly, behind the interparietal; 3.5 pairs of nuchals, 2.5 anterior pairs three times the size of dorsal scale, the other pair twice as the size of dorsal scale; 7 supralabials on both sides, fifth and sixth below center of the eye, sixth largest; 2 loreals; 2 preoculars, the lower one much larger than the upper one; nostril in center of nasal, in contact with the first supralabial, rostral, anterior loreal, and frontonasal; 8 supraciliaries, first largest; 1 enlarged anterior temporal, in contact with sixth and seventh supralabials; 2 posterior temporals, lower one overlapping upper one; 7 infralabials, first two in contact with postmental, fifth longest; 3 pairs of chin shields, first pair medially in contact with each other; tympanum deeply sunk; ear with weak auricular lobules on the anterior margin. Dorsal scales smooth, not larger than lateral and ventral scales, eight rows on the back between dorsolateral bands; 30 midbody scale rows; 66 paravertebral scales; ventral scales smooth, in 64 rows; subcaudal scales on the original part of tail not enlarged; 13 smooth lamellae beneath finger IV and 20 beneath toe IV; 2 enlarged precloacal scales, left scale overlapping right one. Hemipenis. Fully everted hemipenis short, smooth, and unforked; clear sulcus spermaticus starting from base to the single lobe; a prominent longitudinal fold along with the sulcus at the proximal part (Fig. 2 E–G). Coloration. In life, overall dorsal coloration dark brown with faint black dots on dorsum; dorsolateral band black with bright spots; lower part of check, body, limbs, and tail yellow to orange; lower part of head and neck white. Free margin of upper add lower eyelids orange to yellow. Eyes with black round pupil and yellow iris. In preservation, color fades; orange and yellow faded to cream or white; overall dorsal and lateral coloration black to dark brown; venter cream. Variation. Paratype ITBCZ 6535 has supraoculars 3 and 4 incompletely fused on both sides, separated by a suture that is about one third of their width. Supraoculars 1 and 2 incompletely separated from each other on right side. Paratype 6536 with prefrontals separated from each other. Table 3 summarizes variation in size and scalation of the type series. Field notes. All specimens were collected at night, on the ground among rotting leaves in secondary forest connected with a cashew plantation. However, the skink was observed being active in the daytime and it may be diurnal. Sexual dimorphism. Males have lower part of cheek, body, limbs, and tail yellow to orange while these parts in female are cream. Fingers and toes not touching in female when limbs adpressed. Distribution. The new species is currently known only from Ba Ra Mountain, Binh Phuoc Province, southern Vietnam (Fig. 4). Based on the continuous habitat and elevation between Ba Ra Mountain and its vicinity, this skink may also occur in Binh Phuoc and Dak Nong provinces (Vietnam) and Mondulkiri Province in eastern Cambodia. Etymology. The specific epithet baraensis is a toponym derived from the Ba Ra Mountain where the new species was discovered. We recommend “Bara ground skink”, “Thằn lằn cổ bà rá”, and “Rắn mối bà rá” as the common English, Vietnamese, and local names of the new species, respectively. Comparisons. Scincella baraensis sp. nov. differs morphologically from its Asian congeners as follows: from S. apraefrontalis by having more midbody scale rows (30 vs. 18), dorsal scales not enlarged (vs. enlarged—distinctly larger than lateral scales [Smith 1935]), more lamellae beneath toe IV (18–20 vs. 8–9), and presence (vs. absence) of prefrontal (Nguyen et al. 2010a); from S. badenensis by having fewer midbody scale rows (30 vs. 32–36), more nuchal pairs (3–3.5 vs. 0–1), fewer temporals (1+2 vs. 2+3), and unforked (vs. forked) hemipenis (Nguyen et a. 2019); from S. barbouri (Stejneger) by having more midbody scale rows (30 vs. 26–28), more lamellae beneath toe IV (18–20 vs. 15–17), and presence (vs. absence) of weak auricular lobules (Stejneger 1925; Ouboter 1986); from S. boettgeri (van Denburgh) by having more lamellae beneath toe IV (18–20 vs. 15–16) and presence (vs. absence) of weak auricular lobules (van Denburgh 1912; Ouboter 1986); from S. capitanea Ouboter by having more lamellae beneath toe IV (18–20 vs. 15–17) and presence (vs. absence) of weak auricular lobules (Ouboter 1986); from S. darevskii by having more midbody scale rows (30 vs. 28), more paravertebral scale rows (66–70 vs. 62), and fewer supraoculars (4 vs. 5) (Nguyen et al. 2010); from S. devorator by having small (vs. enlarged) dorsal scales and presence (vs. absence) of weak auricular lobules (Darevsky et al. 2004; Nguyen et al. 2011); from S. doriae by having small (vs. enlarged) dorsal scales, more dorsal scale rows on back (8 vs. 6), presence (vs. absence) of weak auricular lobules, and absence (vs. presence) of transversely enlarged subcaudals (Boulenger 1887; Smith 1935; Taylor 1963; Bourret 2009); from S. formosensis (van Denburgh) by having by having more lamellae beneath toe IV (18–20 vs. 15–17) and presence (vs. absence) of weak auricular lobules (van Denburgh 1912; Ouboter 1986); from S. huanrenensis Zhao & Huang by having more midbody scale rows (30 vs. 26–28), more lamellae beneath toe IV (18–20 vs. 13–16), and fewer ventral scale rows (64–66 vs. 75–89) (Zhao & Huang 1982; Chen et al. 2001); from S. macrotis (Steindachner) by having small (vs. enlarged) dorsal scales, presence (vs. absence) of weak auricular lobules, and absence (vs. presence) of transversely enlarged subcaudals (Smith 1935); from S. melanosticta by having nuchal scales (3 or 3.5 pair vs. 0), fewer midbody scale rows (30 vs. 34–38), and short and unforked (vs. long and deeply forked) hemipenis (Smith 1935; Taylor 1963; Bourret 2009; Neang et al. 2018); from S. modesta (Gunther) by having small (vs. enlarged) dorsal scales, more lamellae beneath toe IV (18–20 vs. 10–15), and presence (vs. absence) of weak auricular lobules (Smith 1935); from S. monticola by having more midbody scale rows (30 vs. 22–26), more lamellae beneath toe IV (18–20 vs. 10–13), more paravertebrals and ventrals (66–70 and 64–66 vs. 52–59 and 52–58, respectively), and presence (vs. absence) of weak auricular lobules (Schmidt 1927; Neang et al. 2018); from S. nigrofasciata Neang, Chan & Poyarkov by having fewer midbody scale rows (30 vs. 32–33), more nuchal scales (3 or 3.5 pair vs. 0 or 1), more lamellae beneath toe IV (18–20 vs. 15–17), short and unforked (vs. long and deeply forked) hemipenis, and fewer enlarged anterior temporal (1 vs. 2) (Neang et al. 2018); from S. ochracea by having fewer enlarged anterior temporal (1 vs. 2), more lamellae beneath finger IV (12 or 13 vs. 9–11), more dorsal scale rows between dorsolateral lines (8 vs. 6), and absence (vs. presence) of dark vertebral stripe (Bourret 2009; Pham et al. 2015; Neang et al. 2018); from S. potanini (Gunther) by having more midbody scale rows (30 vs. 27), more lamellae beneath toe IV (18–20 vs. 17), and presence (vs. absence) of weak auricular lobules (Gunther 1896); from S. przewalskii (Bedriaga) by having fewer midbody scale rows (30 vs. 22–34), more lamellae beneath toe IV (18–20 vs. 17), more supralabials (7 vs. 6), and more dorsal scale rows between dorsolateral lines (8 vs. 6) (Wang & Zhao 1986); from S. punctatolineata (Boulenger) by having more midbody scale rows (30 vs. 24–26), more lamellae beneath toe IV (18–20 vs. 12–14), and presence (vs. absence) of nuchals (Smith 1935); from S. rara by having small (vs. enlarged) dorsal scales, more midbody scale rows (30 vs. 24), and a single (vs. double) row of lamellae beneath toes and figures II–IV (Darevsky & Orlov 1997); from S. reevesii by having more nuchals (3 or 3.5 pairs vs. 0 or 1), presence (vs. absence) of weak auricular lobules, short and unforked (vs. long and deeply forked) hemipenis, and absence (vs. presence) of slightly enlarged transversal subcaudals (Smith 1935; Bourret 2009; Neang et al. 2018); from S. rufocaudata by having short and unforked (vs. long and deeply forked) hemipenis, fewer midbody scale rows (30 vs. 32–34), and one (vs. two) anterior temporal (Darevsky & Nguyen 1983; Neang et al. 2018); from S. rupicola by having fewer midbody scale rows (30 vs. 34–36), more nuchals (3 or 3.5 pairs vs. 0 or 1), short and unforked (vs. long and deeply forked) hemipenis, and one (vs. two) anterior temporal (Smith 1935; Taylor 1963; Neang et al. 2018); from S. schmidti (Barbour) by having more midbody scale rows (30 vs. 26), presence (vs. absence) of weak auricular lobules, and more lamellae beneath toe IV (18–20 vs. 11) (Barbour 1927); from S. tsinlingensis (Hu & Zhao) by having more lamellae beneath toe IV (18–20 vs. 11–16), presence (vs. absence) of weak auricular lobules, and more dorsal scale rows between dorsolateral bands (8 vs. 6) (Ouboter 1986; Inger et al. 1990); from S. vandenburghi (Schmidt) by having more midbody scale rows (30 vs. 28) and more lamellae beneath toe IV (18–20 vs. 12) (Schmidt 1927); and from S. victoriana (Shreve) by having more midbody scale rows (30 vs. 26), more lamellae beneath toe IV (18–20 vs. 15), and smooth (vs. keeled) scales on dorsum and tail (Ouboter 1986).

A new skink of the genus Scincella Mittleman, 1950, from Binh Phuoc Province, southern Vietnam is described based on morphological data, including hemipenial characters, and nucleotide sequences from COI. The following morphological characters diagnose Scincella baraensis sp. nov.: relatively small size in adults (snout-vent length up to 49.2 mm); when limbs adpressed, toes do not or just reach fingers; 30 smooth midbody scale rows and dorsal scales not enlarged; 66–70 paravertebral scales; 64–66 ventral scale rows; 7 supralabials, the fifth and sixth below the center of the eye; enlarged temporal scales 1+2; nuchals present; ear with small auricular lobules; 18–20 smooth lamellae beneath toe IV; hemipenis smooth, short, and unforked; dorsum with faint black dots. The new species differs from its congeners by an uncorrected p-distance in COI sequences of at least 16.2%.

Acanthosaura phongdienensis sp. nov. Holotype (Fig. 2). KIZ 10695, an adult male collected on 22 October 2010, 16 o 28��� 03.6��� N, 107 o 16��� 40.8��� E, at Phong Dien Nature Reserve, Thua Thien-Hue Province, Vietnam (site 8, Fig. 1), about 197 m a.s.l. by Robert W. Murphy, Wei-Wei Zhou, Ba Dinh Vo, Luan Thanh Nguyen, and Sang Ngoc Nguyen. Paratypes. ROM 48013 and ITBCZ 6830, adult males; KIZ 10657, KIZ 10697, and ITBCZ 6831, adult females; and KIZ 10617, a sub-adult male (Table 3). Other data were the same as for the holotype except for KIZ 10464, which was collected in September 2010 at an elevation of 1019 m a.s.l. Diagnosis. Medium size, SVL 73���77 mm in males and 59���65 mm in females; TL much longer than SVL; rostral scute entire; nuchal spine at middle of nuchal crest and tympanum; tympanum naked; postorbital spine about one half of ED; nuchal and dorsal crests continuous (Figs. 2���4); lateral scales small, intermixed with large, keeled scales whose keels point backward and back-upward; SL 9���12; IL 10���11. Description of holotype. Adult male, SVL 74.2 mm. SL 11, IL 10. Head moderately tall (HH/HL = 0.63) and with HH/HW = 0.94. Rostral entire and wide, nearly trapezoidal (RH/RW = 0.5), upper wider than lower, not touching nostril, bordered posteriorly by four smaller smooth scale and laterally by first supralabials. When mouth closed, rostral in contact with mental and first infralabials. Nasals with pits, not touching supralabials. Scales on dorsum of head large and keeled. Canthus-rostralis-supraciliary scales slightly serrated, composed of 11 enlarged scales, separated from postorbital spine by a notch. Postorbital spine length about one half ED. An enlarged scale row below orbit. Interorbital scales keeled. Nuchal spine present midway between nuchal crest and tympanum, surrounded by spine-like scales. Tympanum naked, not round, surrounded by small scales. TD smaller than ED. A row of large and keeled scales from posterior edge of orbit to nuchal spine. Mental nearly pentagonal, posteriorly in contact with two elongate postmentals that contact with each other. Throat with two large scale rows parallel with infralabials. Gular scales strongly keeled except for smooth scales near the mental. Standard counts and measurements provided in Table 3. Body elongate with dorsal and nuchal crest connected by lower crests (Fig. 2B). Nuchal crests longer than dorsal ones (3.81 mm vs. 1.56 mm) that reduce posteriorly. Dorsal scales heterogeneous, small, imbricate, intermixed with large, strongly keeled scales that have points directed backward or back-upward. An oblique fold in each side of neck, in front of shoulder. Ventrals strongly keeled. Limbs slender. Hind limb stretched forwards reaches nostril. Dorsal surface of limbs with large, strongly keeled scales. Narrow subdigital lamellae under fourth finger and toe 14 and 20, respectively. Tail longer than snout���vent length (TL/SVL = 1.8), covered with uniform strongly keeled scales; postcloacal region of tail strongly swollen. In life, dorsum of body green with black lateral blotches. Venter whitish. Large black area in front of shoulder. Green to yellow-green wide band expands from snout through labials, tympanum, and neck to lateral of body. Upper side of head and neck black. Upper part of lateral view of head and neck also black. Black colored area becomes narrow at neck and expands at dorsum forming a diamond-shaped band in dorsal view. Tips of nuchal and postorbital spines, and nuchal crests yellow-green. Iris brown with a round pupil. Elbow and knee with white blotches. Original tail dark brown with white-cream or light green alternating rings. In preservative (Fig. 2), color faded with unchanged pattern. Upper side of body and limbs with alternating turquoise, white, and black blotches. Labials, tympanum, and lateral view of neck white. DNA Barcode. Type series with GenBank Accession No. MK 695205 ��� MK 695210. sepa- Variation. Males significantly larger than females (SVL 74.9 mm versus 61.4 mm; p p, Published as part of Nguyen, Sang Ngoc, Jin, Jie-Qiong, Vo, Ba Dinh, Nguyen, Luan Thanh, Zhou, Wei-Wei, Che, Jing, Murphy, Robert W. & Zhang, Ya-Ping, 2019, A new species of Acanthosaura Gray 1831 (Reptilia: Agamidae) from central Vietnam, pp. 555-565 in Zootaxa 4612 (4) on pages 559-561, DOI: 10.11646/zootaxa.4612.4.7, http://zenodo.org/record/3235904

Microhyla aurantiventris sp. nov. Figs. 3���7; Tables 4, 5. Holotype: ITBCZ 4361, adult male, collected while calling on the bank of a small pool that formed after heavy rain nearby Ho Chi Minh Tay Road, Tram Lap Forest, Kon Von II Village, Dak Rong Commune, K���Bang District, Gia Lai Province (14.53994��N, 108.45104��E, ca. 1210 m a.s.l., Fig. 9); collected on 31 July 2016 by Luan T. Nguyen, Tam A. Nguyen, Tiep T. Nguyen, and Vy H. Tran. Paratypes: Total 15 specimens, all collected by Luan T. Nguyen, Tam A. Nguyen, Tiep T. Nguyen, and Vy H. Tran at the same location as holotype: ITBCZ 4288, adult male, collected on 29 July 2016; ITBCZ 4360, adult gravid female, collected on 0 1 August 2016; ITBCZ 4362���4372, eleven adult males, collected on 31 July 2016; ITBCZ 4506���4507, two adult males, collected on 16 November 2016. Diagnosis: Microhyla aurantiventris sp. nov. is characterized by a combination of the following characters: 1) males having SVL 25.2���27.0 mm in 15 individuals, single known female with SVL 30.5 mm; 2) body habitus moderately stocky; 3) head flat, wider than long, snout rounded in dorsal and lateral views, slightly protruding in ventral profile; 4) skin on dorsum and flanks slightly shagreened with numerous tiny tubercles, ventral surfaces smooth; 5) finger I well developed, more than one-half the length of finger II; 6) tips of three outer fingers slightly enlarged, forming weak disks and tips of all toes distinctly dilated into wide disks with narrow peripheral grooves; 7) both finger and toe disks with dorsal median longitudinal grooves; 8) three palmar tubercles (inner and medial palmar tubercles elongated; outer palmar tubercle elongated) and two metatarsal tubercles (inner metatarsal elongated, outer metatarsal rounded); 9) tibiotarsal articulation of adpressed limb reaching slightly beyond the orbit;10) webbing formula: I 1 �����2 II 1 �����2�� III 2���3 1 / 3 IV 3 �����1�� V; 11) dorsum yellowish-brown with dark brown markings, chin and throat yellowish-orange to yellow with small dark brown speckling on its margin, belly bright yellowish-orange to golden-yellow; and 12) an advertisement call consisting of 15���26 pulses with a dominant frequency of 1.1���2.2 kHz (as recorded at 18.5��C). The interspecific genetic p -distances from 12S���16S rRNA gene fragment between the new species and its congeners vary from 9.0% to 14.8%. Description of holotype (Figs. 4, 5). Medium-sized specimen, with SVL 26.1 mm; habitus moderately stocky, head length shorter than head width (HL/HW 0.83); snout rounded, slightly prominent in venral profile, longer than diameter of eye (SL/EL 1.25); eye small, slightly protuberant in dorsal view, pupil rounded (Fig. 5, F); dorsal surface of head flat, canthus rostralis distinct, rounded; loreal region weekly concave; nostril rounded, laterally, closer to tip of snout than to eye; tympanum hidden, supratympanic fold weak, running from posterior corner of eye to arm insertion; vomerine teeth absent, tongue roundly spatulate and free behind, lacking papillae, vocal sac opening larger. Forelimbs comparatively short, about three times shorter than hindlimbs (FLL/HLL 0.32); hand longer than lower arm and shorter over two times than forelimb length (HAL/FLL 0.40); fingers slender, rounded in crosssection, first finger well developed, longer by more than one-half of second finger length (1FLO/2FLO 0.64); relative finger lengths: IColoration of holotype in life (Fig. 5). Dorsal surface of head and trunk yellowish-brown; a distinct brown interorbital bar between eyelids, forming a reverse triangle in shape running posteriorly towards scapular region and not covering dorsal surfaces of upper eyelids; a dorsal dark-brown marking consisting of two reverse V-shaped figures forming a hourglass-shaped marking: anterior reverse V-shaped figure runs from scapular area posteriorly and laterally, opening at level of axilla and getting narrower again posteriorly on middle of dorsum; posterior reverse V-shaped figure starts at middle of dorsum and runs laterally towards groin (Fig. 5, A, B); two black scapular spots present above axilla on each side of body; few smaller black spots irregularly scattered on dorsolateral and lateral sides of body with one larger dark spot on each side just above axilla; mid-vertebral line brown; supratympanic fold yellowish; dorsolateral surfaces of trunk and upper arm yellowish-brown with a few small black spots; two narrow dark brown bars on each forelimb on dorsal surfaces of forearm; dorsal surfaces of thigh, tibia, and tarsus yellowish-brown with dark brown cross-bars and half cross-bars alternating on each hindlimb: two cross-bars and two half cross-bars on each thigh, one cross-bar and four half cross-bars on each shank, one dark cross-bar on each tarsus (Fig. 5, A, B); fingers and toes dorsally beige with dark brown cross-bars; throat and chest dark golden-yellow with dark speckling on margin of throat, ventral skin dark orange-yellow (Fig. 5, C), limbs ventrally yellow with small black spots; hand and foot ventrally reddish brown (Fig. 5, D, E); pupil black, fine golden reticulations throughout iris except for darker area at anterior corner of iris (Fig. 5, F). * holotype * holotype Coloration of holotype in preservative. After preservation in ethanol, dorsal coloration changed to light greyish-brown (Fig. 4, A, C), ventral surface of chest, belly, and limbs whitish-beige (Fig. 4, B); dorsal pattern, dark spots on dorsum and stripes on dorsal surfaces of limbs unchanged, dark brown pattern changed to lighter brown; iris completely black. Variation (Table 4, Fig. 6). Specimens vary in body size, coloration of dorsal surface, form of dark brown markings on the dorsum, and in the coloration of ventral surfaces: the single gravid female paratype (ITBCZ 4360) has a larger body size than male paratypes (SVL 30.5 mm vs. 26.15�� 0.52 mm [25.2���27.0; n=15] in males); males ITBCZ 4365 and ITBCZ 4369 have a much darker dorsal coloration���a dark brown background color with dark blackish-brown markings, male ITBCZ 4365 has less distinct posterior dark brown reverse V-shaped figure on dorsum (see Fig. 6); males ITBCZ 4364���4365 and ITBCZ 4368���4369 have more dark spots in the axilla area compared to the holotype. The female paratype ITBCZ 4360 has a lighter yellowish ventral surface (vs. orange to yellowish-orange in males, see Fig. 6, lower part) and a yellowish-beige throat with brown speckling. Two paratypes (ITBCZ 4506���4507) and four non-vouchered specimens observed on 16 November 2016 have a lighter yellowish ventral coloration compared to males collected in the end of July and in August 2016. Advertisement call. The advertisement call description is based on recordings of paratypes ITBCZ 5406 and ITBCZ 5407; all calls were recorded at 18.5��C ambient temperature (Fig. 7, Table 5). Calls consisted of pulses produced in series, sounding like a quacking duck. Call duration ranged from 110 to 227 ms, and calls were repeated at a rate of 3.1���3.9 calls per second with a 88���391 ms intercall interval. Calls contained from 15 to 26 pulses and had an average pulse rate of 110���144 pulses per second. Dominant frequency varied from 1.8 to 2.2 kHz. Rise-time to reach full amplitude was 67���199 ms, and fall-time from the peak to the end of call varied from 27���48 ms (Table 5). Harmonics were present at approximately 4.3, 6.3, 10.5, 17.3, and 19.5 kHz. No fundamental frequency was detected (Fig. 7). Natural history. All specimens were collected at night from 19:00 to 00:00 h on the banks of a small temporary pond that was formed after heavy rain along the sides of a recently constructed road (Fig. 9). Male frogs were found calling from the small dry branch of a dead tree in close proximity to a water-filled pool. The single female was found about 1 m from a temporary pond at night. Until recently, the survey area was covered by evergreen polydominant montane primary forests. Construction of the Ho Chi Minh Road (completed in 2013) across the area damaged forested areas and facilitated formation of temporary waterbodies along the road embankment. Calling males were also heard at around 15:00 h. The new species was found in sympatry with four congeners including M butleri, M. heymonsi, M. mukhlesuri, and M. pulverata (Fig. 10) and microhylids Kaloula indochiensis Chan, Blackburn, Murphy, Stuart, Emmett, Ho & Brown and Micryletta inornata (Boulenger), all of which were reproducing simultaneously with the new species, sharing the same breeding site. Other anurans such as Feihyla palpebralis (Smith), Feihyla vittata (Boulenger), Polypedates sp. (leucomystax species complex), Rhacophorus kio Ohler & Delorme, R. rhodopus Liu & Hu, Fejervarya limnocharis (Gravenhorst), and Occidozyga martensii (Peters) also occurred in sympatry. Larval stages and eggs of the new species are unknown. Distribution. Microhyla aurantiventris sp. nov. is currently known only from the type locality in Tram Lap Forest, Kon Von II Village, Dak Rong Commune, K���Bang District, Gia Lai Province, Vietnam (Fig. 1). The species was recorded from elevation ca. 1210 m a.s.l. The distribution of the new species is unknown, and discovery of new localities within the Kon Tum Plateau is anticipated. Conservation status. Currently, evergreen forest in Tram Lap connects with other forests in Kon Tum and Gia Lai Provinces. Based on its habitat and distribution elevation, the new species is likely to be endemic to Kon Tum Plateau. However, the extent of its actual distribution range requires further study. Given the available information, we suggest Microhyla aurantiventris sp. nov. be considered as Data Deficient following IUCN���s Red List categories (IUCN Standards and Petitions Subcommittee 2016). Etymology. The specific name ��� aurantiventris ��� is a Latin adjective in the nominative singular, feminine gender, derived from ��� aurantiacus ���������orange-colored��� and ��� venter ���������belly���, referring to the distinctive bright orange-yellow coloration of ventral surfaces in adult males of the new species. The recommended common name in English is ���Orange-bellied narrow-mouth frog���. The recommended common name in Vietnamese is ���Nh��i b���u b���ng v��ng���. Comparisons. In having medium body size for Microhyla (SVL 25.2���27.0 mm in 15 males and 30.5 mm in a single female), M. aurantiventris sp. nov. can be distinguished from the smaller members of the genus including M. achatina (male 16 mm, female 23 mm), M. annamensis (males 15.2���19.8 mm, females 18.2���22.6 mm), M. annectens, M. arboricola Poyarkov, Vassilieva, Orlov, Galoyan, Tran, Le, Kretova & Geissler, M. borneensis, M. chakrapanii Pillai, M. fissipes; M. fusca, M. heymonsi, M. karunaratnei Fernando & Siriwardhane, M. kodial, M. laterite Seshadri, Singal, Priti, Ravikanth, Vidisha, Saurabh, Pratik & Gururaja, M. maculifera Inger, M. malang, M. marmorata, M. minuta Poyarkov, Vassilieva, Orlov, Galoyan, Tran, Le, Kretova & Geissler, M. mixtura, M. mukhlesuri, M. mymensinghensis Hasan, Islam, Kuramoto, Kurabayashi & Sumida, M. nanapollexa, M. nilphamariensis, M. orientalis, M. ornata, M. palmipes, M. perparva, M. petrigena, M. pineticola, M. pulchella, M. pulverata, M. sholigari, M. superciliaris, Microhyla taraiensis, and M. zeylanica (Table 6). * The holotype specimen of M. fusca is male (N.A. Poyarkov, unpublished data). In having moderately stocky body habitus (defined by Bain & Nguyen 2004, Poyarkov et al. 2014), the new species can be distinguished from congeners that have slender or moderately slender bodies, including M. achatina, M. annectens, M. berdmorei, M. butleri, M. fusca, M. mihintalei, M. minuta, M. nanapollexa, M. okinavensis, M. orientalis, M. ornata, M. palmipes, M. sholigari, M. superciliaris, and M. zeylanica. In having first finger well-developed (1FLO>��2FLO), Microhyla aurantiventris sp. nov. can be diagnosed from M. borneensis, M. nanapollexa, M. palmipes, M. perparva, and M. petrigena, which have first finger nub or bulge. In having outer metatarsal tubercle oval-shaped, M. aurantiventris sp. nov. can be separated from the burrowing species M. mihintalei, M. picta Schenkel, and M. rubra, which have enlarged shovel-shaped outer metatarsal tubercles used for digging. In having comparatively short hindlimbs (tibiotarsal articulation of adpressed hindlimb reaching slightly beyond the orbit but not reaching the tip of snout) the new species can be distinguished from M. achatina, M. annamensis, M. annectens, M. arboricola, M. darevskii, M. fissipes, M. mantheyi, M. marmorata, M. mukhlesuri, M. mymensinghensis, M. nanapollexa, M. perparva, M. petrigena, M. pulchella, M. pulchra, M. pulverata, and M. superciliaries, which have tibiotarsal articulation of an adpressed hindlimb reaching just, or well, beyond the snout. In having weak disks on fingers II���IV, well-developed disks on all toes, and median longitudinal grooves on dorsal surfaces of fingers II���IV and all toes, M. aurantiventris sp. nov. differs from the M. ornata species group (M. ornata, M. rubra, M. mihintalei, M. nilphamariensis, and M. taraiensis) and the M. fissipes species group (M. fissipes, M. mukhlesuri, M. mymensinghensis, and M. okinavensis), as well as from M. picta, M. pulchra, and M. zeylanica; these species do not have disks on fingers and toes and lack median longitudinal grooves on dorsal surface of digits. The new species can be diagnosed from M. chakrapanii, M. darevskii, and M. maculifera, which have no disks on fingers and lack median longitudinal grooves on dorsal surface of digits. Microhyla aurantiventris sp. nov. can be further differentiated from M. kodial, M. mixtura, M. sholigari, and M. superciliaries, which have no dorsal median longitudinal grooves on finger disks. The new species differs from M. annectens, M. arboricola, M. maculifera, M. perparva, and M. petrigena of M. annectens species group by having two metatarsal tubercles (vs. one metatarsal tubercle). In having toes with basal webbing (webbing formula I 1 �����2 II 1 �����2�� III 2���3 1 / 3 IV 3 �����1�� V), M. aurantiventris sp. nov. can be distinguished from the following members of its genus that have fully developed webbing reaching to disks at most toes (usually with the exception of toe IV): M. annamensis, M. annectens, M. berdmorei, M. darevskii, M. malang, M. mantheyi, M. marmorata, M. nanapollexa, M. perparva, M. petrigena, M. pulchella, M. pulverata, and M. superciliaris (detailed by Poyarkov et al. 2014). Toe webbing of M. arboricola and M. pulchra reaches the level of distal subarticular tubercles and, thus, they can be diagnosed from the new species. The following species of Microhyla show less developed rudimentary toe-webbing not exceeding the level of proximal subarticular tubercles: M. achatina, M. borneensis, M. fissipes, M. heymonsi, M. karunaratnei, M. minuta, M. mixtura, M. mukhlesuri, M. mymensinghensis, M. okinavensis, M. orientalis, M. ornata, M. palmipes, M. picta, M. pineticola, M. rubra, M. sholigari, and M. zeylanica (Poyarkov et al. 2014). By possessing a weak, discontinuous dorsomedial (vertebral) line, M. aurantiventris sp. nov. can be distinguished from M. achatina, M. heymonsi, M. nilphamariensi s, and M. pineticola, which have a distinct and continuous light dorsomedial line. The new species can be further separated from M. heymonsi and M. pineticola by absence of characteristic ()-shaped black spot in the middle o, Published as part of Nguyen, Luan Thanh, Poyarkov, Nikolay A., Nguyen, Tiep Tan, Nguyen, Tam Ai, Tran, Vy Huu, Gorin, Vladislav A., Murphy, Robert W. & Nguyen, Sang Ngoc, 2019, A new species of the genus Microhyla Tschudi, 1838 (Amphibia: Anura: Microhylidae) from Tay Nguyen Plateau, Central Vietnam, pp. 549-580 in Zootaxa 4543 (4) on pages 562-574, DOI: 10.11646/zootaxa.4543.4.4, http://zenodo.org/record/2618037, {"references":["IUCN Standards and Petitions Subcommittee (2016) Guidelines for Using the IUCN Red List Categories and Criteria. Fersion 12. Prepared by the Standards and Petitions Subcommittee. Available from: http: // www. iucnredlist. org / documents / RedListGuidelines. pdf (accessed 12 March 2017)","Bain, R. H. & Nguyen, Q. T. (2004) Three new species of narrow-mouth frogs (Genus: Microhyla) from Indochina, with comments on Microhyla annamensis and Microhyla palmipes. Copeia, 2004 (3), 507 - 524. https: // doi. org / 10.1643 / ch- 04 - 020 r 2","Poyarkov, J. N. A., Vassilieva, A. B., Orlov, N. L., Galoyan, E. A., Tran, D., Le, D. T. T., Kretova, V. D. & Geissler, P. (2014) Taxonomy and distribution of narrow-mouth frogs of the genus Microhyla Tschudi, 1838 (Anura: Microhylidae) from Vietnam with descriptions of five new species. Russian Journal of Herpetology, 21 (2), 89 ‾ 148.","Yuan, Z. Y., Suwannapoom, C., Yan, F., Poyarkov, N. A., Nguyen, S. N., Chen, H. M., Chomdej, S., Murphy, R. W. & Che, J. (2016) Red River barrier and Pleistocene climatic fuctuations shaped the genetic structure of Microhyla fissipes complex (Anura: Microhylidae) in southern China and Indochina. Current Zoology, 62 (6), 531 - 543. https: // doi. org / 10.1093 / cz / zow 042","Parker, H. W. (1934) Monograph of the frogs of the family Microhylidae. Trustees of the British Museum, London, 212 pp.","Boulenger, G. A. (1900) Descriptions of new batrachians and reptiles from the Larut Hills, Perak. In: J. Hornell (Ed), The Annals and magazine of natural history; zoology, botany, and geology being a continuation of the Annals combined with Loudon and Charlesworth's Magazine of Natural History. Taylor and Francis, Ltd., London, pp. 186 - 193. https: // doi. org / 10.1080 / 00222930008678356","Taylor, E. H. (1962) The amphibian fauna of Thailand. The University of Kansas science bulletin, 43, 265 ‾ 599. https: // doi. org / 10.5962 / bhl. part. 13347","Bourret, R. (1942) Les Batraciens de l'Indochine. Institut Oceanographique de l'Indochine, Ha Noi, x + 547 pp. + 4 pls."]}

Xie, Yulin, Wang, Ping, Zhong, Guanghui, Zhu, Fei, Liu, Qin, Che, Jing, Shi, Lei, Murphy, Robert W., Guo, Peng (2018): Molecular phylogeny found the distribution of Bungarus candidus in China (Squamata: Elapidae). Zoological Systematics 43 (1): 109-117, DOI: 10.11865/zs.201810

Sphenomorphus yersini sp. nov. (Figs. 1–4) Holotype. ITBCZ 5685, adult male, collected from Hon Ba NR., Khanh Hoa Province, Vietnam; coordinates 12°8’13”N, 108°57’39”E; elevation 1162 m a.s.l. by L.T. Nguyen, V.D.H. Nguyen, and S.N. Nguyen on 16 October 2016 (Fig. 1). Paratypes. Two specimens, also collected from Hon Ba NR by L.T. Nguyen, V.D.H. Nguyen, and S.N. Nguyen: ITBCZ 5686, adult female (Fig. 2C&D), and ITBCZ 5684, adult male (Figs. 2A&B and 3), collected on 14 October 2016, coordinates 12°8’22”N, 108°58’06”E; elevation 932 m a.s.l. Referred specimen. HBA 44 (released, Fig. 4), adult female, collected on 19 October 2016, coordinates 12°8’22”N, 108°58’06”E; elevation 932 m a.s.l. Diagnosis. Sphenomorphus yersini sp. nov. is distinguished from all of its congeners by a combination of the following morphological characters: size in adults (SVL) up to 56 mm; TaL/SVL ratio 1.81; toes reach to fingers when limbs adpressed; 32–34 smooth midbody scale rows; 61–69 paravertebral scales; 58–67 ventral scale rows; 112 subcaudal scales; four, rarely five, supraoculars; prefrontals in broad contact with one another; two loreal scales; tympanum deeply sunk; smooth lamellae beneath Finger IV and Toe IV 10–12 and 18 –20, respectively; two enlarged precloacal scales; hemipenis smooth, deeply forked, asymmetrical with a long lobe and another short; black and interruptive dorsolateral line; lateral side and lower part of head, neck, and tail orange to red in male. Description of holotype. Adult male; head small and elongate (HW/HL = 0.7); SVL 54 mm; tail long (TaL/ SVL = 1.81), 98 mm in length; lower eyelid scaly; tympanum deeply sunk with a prominent oblique edge; limbs pentadactyl, toes reach to fingers when limbs adpressed. Head scalation smooth; rostral convex, distinctly visible from above, in broad contact with frontonasal, which is broader than long; no supranasals; prefrontals in broad contact with one another; 5 supraoculars on left side and 4 on right side; a pair of frontoparietals, shorter than frontal; frontal narrowing posteriorly, longer than wide, bordered laterally by first two supraoculars, anteriorly by prefrontals, and posteriorly by frontoparietals; frontoparietals in contact with the second to fifth supraoculars on left side and the second to fourth supraoculars on right side; parietals in contact posteriorly, behind the interparietal; no nuchal; 7 supralabials on both sides, the fifth and sixth below the eye; 2 loreals, posterior loreal larger than anterior; nostril in center of nasal, which is in contact with the first supralabial, rostral, anterior loreal, and frontonasal; 8 supraciliaries, first largest; 1 anterior temporal, in contact with sixth and seventh supralabials; 2 secondary temporals, lower temporal overlapping upper one and in contact with seventh supralabial, upper temporal larger, in contact with parietal; lower eyelid scaly, 2 scales in center larger than scales in posterior and anterior areas; 6 infralabials, first pair medially in contact with each other; 1 anterior and 2 postmentals. Dorsal scales smooth, not larger than lateral and ventral scales; 34 midbody scale rows; 61 paravertebral scales; ventral scales smooth, in 60 rows; 112 subcaudal scales, slightly enlarged posteriorly; 11 and 20 smooth lamellae beneath finger IV and toe IV, respectively; 2 enlarged precloacal scales. Hemipenes. Each hemipenis deeply forked and asymmetrical with two smooth lobes. In fully everted position, the inner lobe is shorter and the outer lobe is much longer. Clear sulcus spermaticus starts from the base, divides into two branches at the fork and extends to the tips of the two lobes; body of the long lobe forming regular transversal shallow grooves and ending with terminal papillae. Short lobe obtuse, not forming regular transversal shallow grooves (Fig. 1F). Coloration. In life, overall dorsal coloration black (Hex #000000) to dark brown (Hex #654321) with two interrupted copper (Hex # B87333) dorsal lines in margins of dorsum extending from neck to base of tail; black and interrupted lateral lines under the copper ones; dorsum with irregular black spots; lateral side and lower part of head, neck, and tail orange (Hex #FF6600) to red (Hex #FE2712); venter from chest to tail base yellow (Hex #FFD300) to cream (Hex # FFFDD 0); upper side of limbs black with bright dots. Lateral margins of supralabials and infralabials with black blotches. Free margin of upper eyelid orange and the margin of lower eyelid yellow (Hex #FFD300). Eyes black. In preservation, color fades; orange and yellow disappear; overall dorsal coloration black with two interrupted brighter lines in margins of dorsum extending from neck to base of tail; venter zebra white (Hex #F5F5F5). Variation. Table 2 summarizes variation in size and scalation of the holotype, paratypes, and referred specimen. Supraoculars four in all specimens except for the left side of the holotype which bears 5 scales. Midbody scale rows vary from 32 to 34. Paravertebral and ventral scale rows range from 61 to 69 and from 58 to 67, respectively. Hemipenes of the second paratype ITBCZ 5684 (Fig. 3) similar to those of the holotype in being deeply forked and asymmetrical. Natural history. All specimens were collected at night and under rotting leaf layer in evergreen forest (Fig. 5), elevations 932 m and 1162 m a.s.l., between 19:00–23:30. However, the species was observed being active in the daytime and it may be diurnal. Sphenomorphus yersini sp. nov. was observed to be sympatric with other congener, S. indicus. Sexual dimorphism. Males are larger than females (SVL 56 mm vs. 52 mm, n=4) and have orange (Hex #FF6600) or red (Hex #FE2712) color on lateral sides and lower part of head, neck, and tail. The color on the lateral side in females faded to yellow (Hex #FFD300) or brown (Hex #964B00) with bright spots; lower part of head, limbs, and tail white (Hex #FFFFFF) to zebra white (Hex #F5F5F5); venter from neck to vent yellow (Hex #FCE883). Distribution. The new species is currently known only from Hon Ba NR, Khanh Hoa Province, southern Vietnam (Fig. 5). Etymology. We name this new species in honor of the famous physician and bacteriologist, Alexandre Yersin (1863–1943), who discovered the bacterium responsible for bubonic plague. Hon Ba NR associates with the name of Alexandre Yersin who built a research station on the top of the mountain and worked there. Currently, the research station has been reconstructed and opened to visitors. We recommend Yersin’s Forest Skink as the common name of this new species. Comparisons. Sphenomorphus yersini sp. nov. differs from its congeners in Indochina (Vietnam, Laos, Cambodia, Thailand, Myanmar, Peninsular Malaysia, and southern China [Yunnan]) as follows: from S. anomalopus (Boulenger) by having a smaller size (SVL 50–56 mm vs. 70 mm), fewer midbody scale rows (32–34 vs. 38) and more lamellae beneath toe IV (18–20 vs. 14); from S. bacboensis by having one (vs. two) anterior temporal, more midbody scale rows (32–34 vs. 30–32), and more supralabials (7 vs. 6); from S. cameronicus Smith by having a smaller size (SVL 50–56 mm vs. 70 mm) and fewer midbody scale rows (32–34 vs. 38); from S. cophias Boulenger by having a larger size (SVL 50–56 mm vs. 37 mm), more midbody scale rows (32–34 vs. 24), more lamellae under fourth toe (18–20 vs. 9) and prefrontals in broad contact (vs. separated); from S. cryptotis by having a smaller size (SVL 50–56 mm vs. 58–79 mm), fewer midbody scale rows (32–34 vs. 36–39) and tympanum deeply sunk (vs. superficial); from S. grandisonae Taylor by having a larger size (SVL 50–56 mm vs. 30 mm), one (vs. two) anterior temporal, more supralabials (7 vs. 6) and lamellae under toe IV (18–20 vs. 12), and adpressed limbs overlapping (vs. failing to touch); from S. helenae Cochran by having more midbody scale rows (32–34 vs. 30), prefrontals in broad contact (vs. separated), and presence of an interrupted (vs. uninterrupted) lateral stripe; from S. incognitus by having a smaller size (SVL 50–56 mm vs. 80–103 mm), fewer midbody scale rows (32–34 vs. 36–40), and one (vs. two) anterior temporal; from S. indicus by having a smaller size (SVL 50–56 mm vs. 90 mm), prefrontals in broad contact (vs. separated), and asymmetrical and deeply forked hemipenis (vs. symmetrical); from S. lineopunctulatus Taylor by having a smaller size (SVL 50–56 mm vs. 84 mm), fewer midbody scale rows (32–34 vs. 38), fewer paravertebral scale rows (61–69 vs. 76), and prefrontals in broad contact (vs. separated); from S. maculatus by having a smaller size (SVL 50–56 mm vs. 62 mm), fewer midbody scale rows (32–34 vs. 38–42), and prefrontals in broad contact (vs. separated); from S. malayanum by having fewer ventral scales (32–34 vs. 74), fewer paravertebral scales (61–69 vs. 76–80), more lamellae under fourth toe (18–20 vs. 15), and deeply sunk (vs. shallow) tympanum; from S. mimicus by having a larger size (SVL 50–56 mm vs. 36 mm), more midbody scale rows (32–34 vs. 30) and more lamellae under fourth toe (18–20 vs. 16); from S. orientale (Shreve) by having more midbody scale rows (32–34 vs. 24–26) and fewer paravertebral scale rows (61– 69 vs. 69–71); and from S. praesignis (Boulenger) by having a smaller size (SVL 50–56 mm vs. 109 mm) and more midbody scale rows (32–34 vs. 28). Sphenomorphus yersini sp. nov. differs from S. sanctus (Duméril & Bibron) by having a larger size (SVL 50–56 mm vs. 40–45 mm), fewer paravertebral scales (61–69 vs. 71), fewer supraoculars (4[5] vs. 5), and fewer lamellae under toe IV (18–20 vs. 26–27); from S. scotophilus (Boulenger) by having more midbody scale rows (32–34 vs. 28–31), fewer supraoculars (4 vs. 5), and fewer lamellae under fourth toe (18–20 vs. 22–23); from S. senja Grismer & Quah by having a smaller size (SVL 50–56 mm vs. 60–65 mm), fewer paravertebral scales (61–69 vs. 72–73), fewer ventral scale rows (60–67 vs. 68), one (vs. two) anterior temporal, more lamellae under toe IV (18–20 vs. 13–17) and prefrontals in broad contact with each other (vs. narrow in contact or slightly separated); from S. sheai by having a larger size (SVL 50–56 mm vs. 35 mm), more midbody scale rows (32–34 vs. 20), more paravertebral scales (61–69 vs. 53), more supralabials (7 vs. 6), more lamellae under fourth toe (18–20 vs. 6), and adpressed limbs overlapping (vs. separated); from S. shelfordi (Boulenger 1900) by having a smaller size (SVL 50–56 mm vs. 67 mm), fewer lamellae under toe IV (18–20 vs. 28–29) and the absence of nuchals (vs. presence of a single pair of nuchals); from S. stellatus by having a smaller size (SVL 50–56 mm vs. 80 mm), more midbody scale rows (32–34 vs. 24) and absence (vs. presence) of two enlarged, broader than long, vertebral scale rows; from S. sungaicolus Sumarli, Grismer, Wood, Ahmad, Rizal, Ismail, Izam, Ahmad & Linkem by having a smaller size (SVL 50–56 mm vs. 67–90 mm), fewer midbody scale rows (32–34 vs. 39–44), fewer paravertebral scales (61–69 vs. 72–81) and fewer ventral scale rows (32–34 vs. 74–86); from S. tarsus (Smith) by having a smaller size (SVL 50–56 mm vs. 90–92 mm) and two loreals (vs. three); from S. tetradactylus by having more midbody scale rows (32–34 vs. 20), absence (vs. presence) of external ear opening, and forelimb with five digits (vs. four digits); from S. tonkinensis by having a larger size (SVL 50–56 mm vs. 36–49 mm), TaL/ SVL ratio 1.80 (vs. 1.32), and one (vs. two) anterior temporal; from S. tridigitus by having a larger size (SVL 50– 56 vs. 35), more midbody scale rows (32–34 vs. 20), absence (vs. presence) of external ear opening and forelimb with five digits (vs. three digits); and from S. tritaeniatus by having a larger size (SVL 50–56 vs. 47), fewer midbody scale rows (32–34 vs. 38) and paravertebral scales (61–69 vs. 81), more lamellae under toe IV (18–20 vs. 15) and one anterior temporal (vs. two). Sphenomorphus yersini sp. nov. is similar to S. buenloicus in many aspects (size, midbody scale rows, number of anterior temporals, lamellae under fourth toe, etc.). However, the new species can be distinguished from S. buenloicus by having a relatively longer tail (TaL/SVL ratio 1.8 vs. 1.2), usually more ventral scale rows (58–67 vs. 55–58) and deeply forked hemipenis with two asymmetrical lobes and the outer lobe is much longer than the inner one (vs. hemipenis forked at the middle point of its length with two developing lobes and the outer lobe is slightly longer than the inner one [Figs. 6 & 7]).

Nguyen, Luan Thanh, Dzung Trung Le, Nikolay A. Poyarkov Jr., Vo, Ba Dinh, Phan, Hoa Thi, Duong, Tang Van, Murphy, Robert W., Nguyen, Sang Ngoc (2018): A new species of Leptolalax (Anura: Megophryidae) from Son Tra Peninsula, central Vietnam. Zootaxa 4388 (1): 1-21, DOI: https://doi.org/10.11646/zootaxa.4388.1.1

Leptolalax rowleyae sp. nov. FIgS. 2���5; TAbLeS 3���5. Holotype. ITBCZ 2783, ADULT MALe, COLLeCTeD WHILe CALLINg fROM A SMALL HOLe UNDeR A ROCK NeAR THe HeADWATeR Of A SMALL STReAM WITH 1.5��� 2.0 M IN WIDTH IN eVeRgReeN TROPICAL fOReST IN SON TRA NR, DA NANg CITY, VIeTNAM (16.12754��N, 108.24634��E, CA. 380 M A.S.L.); COLLeCTeD ON 0 8 JANUARY 2013 bY LUAN T. NgUYeN. Paratypes. FOURTeeN SPeCIMeNS, ALL COLLeCTeD fROM SON TRA NR, DA NANg CITY, VIeTNAM: ITBCZ 4502, A gRAVID feMALe, ReCORDeD ON fOReST fLOOR, AbOUT 5 M IN DISTANCe fROM A SMALL STReAM (16.12248��N, 108.29821��E, CA. 450 M A.S.L.), COLLeCTeD ON 10 AUgUST 2013 bY LUAN T. NgUYeN; ITBCZ 2779���81, THRee ADULT MALeS (ALL COLLeCTeD WHILe CALLINg), ReCORDeD NeAR A CASCADe STReAM IN TROPICAL eVeRgReeN fOReST (16.13538��N, 108.25198��E, CA. 440 M A.S.L.), COLLeCTeD ON 0 3 JANUARY 2012 bY LUAN T. NgUYeN AND HOA T. PHAN; ITBCZ 2782, ITBCZ 2784���9, SeVeN ADULT MALeS (ALL ObSeRVeD AND COLLeCTeD WHILe CALLINg), COLLeCTeD ON 0 9 JANUARY 2013 AT THe SAMe LOCATION AS THe HOLOTYPe bY LUAN T. NgUYeN; ITBCZ 4471, A gRAVID feMALe AND ITBCZ 4472���3, TWO jUVeNILeS, ReCORDeD ON fOReST fLOOR NeAR A SMALL STReAM (CA. IN 20 M fROM THe STReAM) (16.13534��N, 108.25222��E, CA. 640 M A.S.L.), COLLeCTeD ON 11 SePTeMbeR 2013 bY LUAN T. NgUYeN AND MANH V. Le. Diagnosis. Leptolalax rowleyae sp. nov. IS DISTINgUISHeD fROM ALL Of ITS CONgeNeRS bY A COMbINATION Of THe fOLLOWINg MORPHOLOgICAL CHARACTeRS: (1) ADULT SVL 23.4���25.4 MM IN MALeS AND 27.0���27.8 MM IN feMALeS; (2) PReSeNCe Of DISTINCT DARK/bROWN DORSOLATeRAL MARKINgS WITH bLACK SPOTS ON fLANKS; (3) PINKISH MILK-WHITe TO LIgHTLY bROWN CHeST AND beLLY WITH NUMeROUS WHITe SPeCKLeS; (4) TYMPANUM eXTeRNALLY DISTINCT; (5) NO WebbINg AND LATeRAL DeRMAL fRINgeS ON fINgeRS AND TOeS; (6) PeCTORAL gLANDS COMPARATIVeLY SMALL, COMPRISINg 2.6���3.7% Of SVL IN ADULT MALeS; (7) VeNTROLATeRAL gLANDS INDISTINCT; (8) IRIS bICOLOReD WITH COPPeR-ORANge UPPeR HALf, fADINg TO gOLDeN IN ITS LOWeR THIRD. THe ADVeRTISeMeNT CALL Of THe NeW SPeCIeS, CONSISTINg Of 4���6 NOTeS, LACKINg A DISTINCT INTRODUCTORY NOTe, WITH AN AVeRAge DOMINANT fReqUeNCY Of 3.2���3.5 KHZ, ALSO DISTINgUISHeS THe NeW SPeCIeS fROM Leptolalax SPeCIeS fOR WHICH CALLS ARe KNOWN. THe NeW SPeCIeS IS ALSO MARKeDLY DISTINCT fROM ALL CONgeNeRS fOR WHICH COMPARAbLe SeqUeNCeS ARe AVAILAbLe (MITOCHONDRIAL geNe 16S RRNA; UNCORReCTeD geNeTIC DISTANCe ��7.4%). Description of holotype. SMALL-SIZeD SPeCIMeN IN gOOD STATe Of PReSeRVATION; HAbITUS STOCKY (FIg. 2). VeNTRAL SURfACe Of HOLOTYPe DISSeCTeD ON RIgHT SIDe beLLY, DISSeCTION-LeNgTH CA. 2.0 MM (FIg. 2B). Head. HeAD WIDTH SLIgHTLY SHORTeR THAN HeAD LeNgTH (HDW/HDL 0.91), fLATTeNeD; SNOUT SLIgHTLY TRUNCATe IN DORSAL VIeW AND ROUNDeD IN PROfILe, PROjeCTINg SLIgHTLY beYOND MARgIN Of LOWeR jAW; NOSTRIL OVAL-SHAPeD, SLIgHTLY ROUNDeD, LOCATeD MUCH CLOSeR TO SNOUT THAN TO eYe; CANTHUS ROSTRALIS INDISTINCT, geNTLY ROUNDeD; LOReAL RegION CONCAVe, geNTLY SLOPINg; PUPIL VeRTICAL; eYe DIAMeTeR NOTAbLY SMALLeR THAN SNOUT LeNgTH (EYE/SNT 0.66); TYMPANUM DISTINCT, ROUNDeD, TYMPANUM DIAMeTeR SMALLeR THAN THAT Of eYe; TYMPANIC RIM eLeVATeD ReLATIVe TO SKIN Of TeMPORAL RegION; VOMeRINe TeeTH AbSeNT; PINeAL OCeLLUS AbSeNT; VOCAL SAC OPeNINgS, OVAL, LOCATeD LATeRALLY ON fLOOR Of MOUTH; TONgUe WIDe WITH bROAD, SHALLOW NOTCH AT POSTeRIOR TIP; SUPRATYMPANIC fOLD fORMINg A DISTINCT WIDe RIDge, RUNNINg fROM POSTeRIOR CORNeR Of eYe POSTeRIORLY TOWARDS DORSAL eDge Of TYMPANUM, geNTLY CURVINg DOWNWARDS TOWARDS AXILLA; SUPRATYMPANIC RIDge COMPARATIVeLY SMOOTH WITH feW fLAT TUbeRCLeS (FIg. 2A���C). Forelimbs. FOReLIMbS THIN, SLeNDeR; fINgeR TIPS ROUNDeD, VeRY SLIgHTLY SWOLLeN; ReLATIVe fINgeR LeNgTHS: I Hindlimbs. HINDLIMbS SLeNDeR, TIbIA NeARLY HALf Of SNOUT-VeNT LeNgTH (TIB/ SVL 0.49). TOe TIPS SIMILAR THOSe Of fINgeRS; ReLATIVe TOe LeNgTHS: I Skin texture and skin glands. SKIN ON DORSUM MOSTLY SMOOTH WITH NUMeROUS TINY TUbeRCLeS AND PUSTULeS fINeLY AND ReLATIVeLY eVeNLY SCATTeReD ON DORSAL AND LATeRAL SURfACeS Of TRUNK, HeAD AND LIMbS; UPPeR eYeLID WITH NUMeROUS SMALL TUbeRCLeS; SKIN ON VeNTRAL SURfACeS Of TRUNK, HeAD AND LIMbS COMPLeTeLY SMOOTH; PeCTORAL gLANDS LOCATeD AT fOReLIMb bASIS ON VeNTRAL SURfACe Of AXILLARY RegION, ROUNDeD, 0.8 MM IN DIAMeTeR, DISTINCT IN LIfe AND IN PReSeRVATIVe; feMORAL gLANDS SMALL, OVAL, APPROXIMATeLY 0.9 MM IN DIAMeTeR, LOCATeD ON POSTeROVeNTRAL SURfACeS Of THIgHS, CLOSeR TO KNee THAN TO VeNT, DISTINCT IN LIfe AND IN PReSeRVATIVe; SUPRA-AXILLARY gLAND LOCATeD IN AXILLARY RegION DORSALLY fROM INSeRTION Of fOReLIMb, OVAL, fLATTeD, 1.1 MM IN DIAMeTeR. VeNTROLATeRAL gLANDS INDISTINCT, ROUNDeD, fLAT, fORMINg AN INDISTINCT LINe (FIg. 2A���C). Color of holotype in life. COLORATION Of HOLOTYPe ReCORDeD AT NIgHT jUST AfTeR CAPTURe (FIg. 2F,G). DORSAL SURfACe Of TRUNK LIgHT bROWN; ONe V-SHAPeD INTeRORbITAL bAR MARKINg WITH PALe CReAM eDgINg ANTeRIORLY; SNOUT DORSALLY beIge; DARK-bROWN W-SHAPeD MARKINg AT SCAPULAR RegION WITH INDISTINCT LIgHT-bROWN eDgINg. DARK bLACKISH-bROWN LINe WITH UNCLeAR bORDeRS LATeRALLY ALONg CANTHUS ROSTRALIS, eNCOMPASSINg NAReS RUNNINg TOWARDS ANTeRIOR CORNeR Of eYe AND A LIgHTeR bROWN LINe MeDIALLY ALONg CANTHUS ROSTRALIS RUNNINg TOWARDS UPPeR eYeLID; DISTINCT bLACKISH SPOT IN ANTeRIOR CORNeR Of eYe; DARK bLACKISH STRIPe beLOW SUPRATYMPANIC RIDge RUNNINg fROM POSTeRIOR CORNeR Of eYe TOWARDS TYMPANUM, TeRMINATINg AbOVe AXILLA, DARK LINe eNCOMPASSINg MOST Of TYMPANUM fORMINg A CHARACTeRISTIC DIAMOND-SHAPeD DARK MARKINg IN TeMPORAL RegION; UPPeR eYeLID LIgHT bROWN WITH SMALL WHITISH SPOTS ON CILIARY eDge; CANTHUS ROSTRALIS ReDDISH-bROWN; DORSAL SURfACeS Of fOReLIMbS (LOWeR ARMS) AND HINDLIMbS LIgHTeR bROWNISH, fOReARMS, AS WeLL AS DORSAL SURfACeS Of fINgeRS AND TOeS MUCH LIgHTeR WITH ORANge bACKgROUND COLOR AND SMALL ReDDISH-bROWN TUbeRCLeS; TRANSVeRSe DARK-bROWN bARS ON DORSAL SURfACe Of THIgHS, TIbIA, TARSUS, LOWeR ARMS, fINgeRS AND TOeS; WHeN HINDLIMb IS fLeXeD, TRANSVeRSe DARK bARS ON DORSAL SURfACeS Of THIgHS CONTINUe ON DORSAL SURfACeS Of SHANKS AND TARSUS (FIg. 2F); eLbOWS AND UPPeR ARMS ORANge WITH LIgHT ReDDISH-bROWN SPOTS ON TUbeRCLeS; LATeRAL SURfACeS Of TRUNK LIgHT PURPLISH-bROWN; VeNTRAL SURfACeS LIgHTLY COLOReD, PINKISH MILK-WHITe WITH DeNSe WHITISH SPeCKLINg eVeNLY SCATTeReD ON eNTIRe VeNTRAL SURfACe INCLUDINg THROAT, CHeST, ARMS, LegS, AND jAWS; WHITISH fLeCKS geTTINg LARgeR AND eSPeCIALLY NUMeROUS ON LATeRAL SIDeS Of bODY. TIP Of fINgeRS AND TOeS ARe WHITISH-PURPLe. LATeRAL SIDeS Of bODY WITH 7 DISTINCT bLACKISH-bROWN bLOTCHeS ON eACH SIDe (0.3���1.4 MM IN DIAMeTeR) LOCATeD fROM AXILLA TO gROIN, MOST POSTeRIOR bLACK SPOT LOCATeD IN gROIN AReA IS AbOUT 4.5 TIMeS LARgeR THAN OTHeR SPOTS. THRee DISTINCT bLACKISH-gReY SPOTS ON LIgHT bROWN bACKgROUND COLOR ON eDge Of UPPeR jAW LOCATeD fROM SNOUT TO AReA UNDeR eYe; MIDDLe ONe LASTINg DORSALLY TOWARDS NARIS eNCOMPASSINg MOST PART Of LOReAL AReA (FIg. 2F). SUPRA-AXILLARY gLAND MILK-WHITe; feMORAL, PeCTORAL, AND VeNTROLATeRAL gLANDS WHITe. IRIS bICOLOReD: COPPeR-ORANge IN UPPeR HALf, fADINg TO gOLDeN IN LOWeR THIRD; fINe bLACK ReTICULATIONS THROUgHOUT IRIS. Color of holotype in preservative. IN PReSeRVATION COLORATION fADeS TO DARK gReY-bROWN ON DORSUM AND fLANKS (FIg. 2A���E); IRIS COLORATION fADeS TO COMPLeTeLY DARK bROWN. BANDINg ON LIMbS fADe TO bROWNISH COLOR AND TURN LeSS DISTINCT; WHITe fLeCKS ON bODY fLANKS AND VeNTRAL SIDeS ReMAIN PRONOUNCeD IN PReSeRVATIVe. VeNTRAL SURfACe Of CHeST, beLLY, THROAT, VeNTRAL PORTIONS Of ARMS AND THIgHS fADe TO WHITISH-bROWN; MACROgRANDS TURN WHITISH. Variation. MeASUReMeNTS Of THe TYPe SeRIeS ARe SHOWN IN TAbLe 2 AND RePReSeNTATIVe PHOTOgRAPHS Of PARATYPeS IN LIfe ARe SHOWN IN FIg. 3. SPeCIMeNS VARY IN bODY SIZe, NUMbeR AND SIZe Of bLACK VeNTROLATeRAL bLOTCHeS AND SLIgHTLY IN COLOR AND PATTeRN IN LIfe. MALeS ITBCZ 2779���80 HAVe DARKeR bROWN COLORATION Of DORSAL SURfACeS AND LIgHT-bROWN VeNTRAL SURfACe WITH DeNSe WHITISH SPeCKLINg COMPAReD WITH THe HOLOTYPe. MALe ITBCZ 2787 HAS INDISTINCT DARK V-SHAPeD INTeRORbITAL bAR; MALe ITBCZ 2782 HAS LeSS DISTINCT DARK bARS ON LIMbS, WHILe MALe ITBCZ 2788 HAS VeRY THIN ALMOST INDISTINCT DARK bARS ON LIMbS. MALe ITBCZ 2784 HAS LIgHT-bROWNISH DORSAL SURfACe WITH TWO bIg PALe MARKINgS beTWeeN AXILLAe AND ReDDISH-bROWN TINT ON bODY fLANKS. MALeS ITBCZ 2780���83 HAVe WHITISH-bROWN COLORATION ON VeNTRAL SURfACeS. FeMALeS TeND TO HAVe LARgeR bODY SIZe (SVL 27.0���27.8; N =2) THAN MALeS (SVL 23.4���25.4; N =11). JUVeNILe ITBCZ 4473 HAS LIgHTeR DORSAL COLORATION COMPAReD TO HOLOTYPe. AS IN MANY OTHeR SPeCIeS Of Leptolalax, COLORATION Of DORSAL SURfACeS IS LIgHTeR AT NIgHT AND geTS DARKeR IN DAYTIMe (IN ALL SPeCIMeNS), WHILe VeNTRAL COLORATION Of ALL SPeCIMeNS beCOMeS DARKeR AND CHANgeS COLOR fROM WHITISH TO ReDDISHbROWN OR bROWN AfTeR CAPTURe IN DAYTIMe OR AfTeR eUTHANIZATION. SKIN TeXTURe APPeARS TO be LeSS TUbeRCULATe IN PReSeRVATIVe THAN IT IS IN LIfe. MALeS Of L. rowleyae sp. nov. WHeN CALLINg INfLATe A COMPARATIVeLY LARge SINgLe SUbgULAR VOCAL SAC (FIg. 3C). Advertisement call. AS IN OTHeR MeMbeRS Of L. applebyi SPeCIeS gROUP, THe ADVeRTISeMeNT CALL Of THe NeW SPeCIeS RePReSeNTS A SeRIeS Of fAINT, qUITe RASPINg CLICKINg SIgNALS, TO HUMAN eAR SUPeRfICIALLY ReSeMbLINg A CALL Of AN ORTHOPTeRAN. CALL DeSCRIPTION IS bASeD ON THe ReCORDINg Of THe ADVeRTISeMeNT CALL Of THe HOLOTYPe, TAKeN AT 21.5 ��C AMbIeNT TeMPeRATURe. THe NeW SPeCIeS HAS SINgLe TYPe Of ADVeRTISeMeNT CALLS (N =10) CONSISTeD Of 4���6 NOTeS (MeAN 5.1 NOTeS PeR CALL; N =51), OR CLICKS, Of VARIAbLe (276���435 MS, MeAN 363 MS) DURATION, RePeATeD AT A RATe Of 8.6���11.3 (MeAN 11.2) NOTeS PeR SeCOND (FIg. 4). A DISTINCT INTRODUCTORY NOTe WAS NOT DISCeRNIbLe. NOTe DURATION VARIeD fROM 4���96 MS (MeAN 20 MS; N =51) AND CONSISTeD Of 1���7 DISTINCT PLUSeS (MeAN 2.1 PULSeS PeR NOTe; N =51). THe DOMINANT fReqUeNCY WAS 3.3 KHZ, HARMONICS WeRe NOT CLeAR bUT CAN be DeTeCTeD AT APPROXIMATeLY 10.5 AND 13.8 KHZ, AND THe fUNDAMeNTAL fReqUeNCY WAS NOT eVIDeNT. CALLS HAD AN AVeRAge INTeRCALL INTeRVAL Of 179 MS (68���428 MS; N =10). NOTe INTeRVAL AND THe NUMbeR Of NOTeS PeR CALL VARIeD SLIgHTLY WITHIN CALLINg bOUTS AND AMONg INDIVIDUALS. TAbLe 4 SHOWS THe VARIATION Of OTHeR CALL CHARACTeRISTICS AMONg eXAMINeD INDIVIDUALS Of THe NeW SPeCIeS. Tadpole description. DeSCRIPTION Of LARVAL MORPHOLOgY IS bASeD ON fOUR TADPOLeS ITBCZ 2790, ITBCZ 3579��� 81; GOSNeR (1960) STAgeS 27���38 (FIg. 5). TADPOLeS WeRe COLLeCTeD IN A MOUNTAIN STReAM, WHILe HIDINg beTWeeN PebbLeS ON THe STReAMbeD. TADPOLeS Of THe NeW SPeCIeS WeRe ACTIVe AT NIgHT AND AVOIDeD LIgHT bY HIDINg THeMSeLVeS beTWeeN ROCKS OR DeAD LeAVeS. Dorsal view. BODY Of TADPOLe ReACHeS A LeNgTH Of 13.8 MM IN THe STAge 31; bODY eLLIPTICAL, eLONgATeD (BW/BL 0.55���0.87, MeAN 0.69; N =4), WITH A ROUNDeD SNOUT. NAReS ANTeRODORSALLY POSITIONeD, NeAReR TO TIP Of SNOUT THAN TO PUPIL (RND/NPD 0.32���0.53; IND /IP 0.73���0.93). EYeS DORSOLATeRAL, LOCATeD AT THe POSTeRIOR eDge Of THe ANTeRIOR ONe fOURTH Of bODY LeNgTH. EYeS COMPARATIVeLY SMALL AND DISTINCT (ED/BL 0.06���0.12, MeAN 0.08; IP/BW 0.46���0.51, MeAN 0.48; N =4). THe MAXIMUM WIDTH Of TAIL MUSCULATURe COMPRISeS LeSS THAN A HALf Of bODY WIDTH (TMW/BW 0.42���0.46, MeAN 0.44; N =4) (FIg. 5B). NARIAL APeRTUReS WITH CHARACTeRISTIC geNTLY ROUNDeD RIMS, SLIgHTLY LIfTeD AbOVe bODY SURfACe. Lateral view. BODY LATeRALLY DePReSSeD (BH/BW 0.76���0.83, MeAN 0.79; N =4). CONe-SHAPeD SPIRACLe SINISTRAL, DIReCTeD POSTeRODORSALLY, POSITIONeD LATeRALLY AT POSTeRIOR HALf Of bODY (SS/BL 0.53���0.71, MeAN 0.61; N =4); CONICAL TUbe Of SPIRACLe IS fUSeD TO bODY WITH A fRee SHORT DISTAL PORTION. VeNT TUbe DeXTRAL, SePARATeD fROM bODY AND ATTACHeD TO LOWeR fIN (FIg. 5A). Tail. TAIL MUSCULAR, LONg, TAIL LeNgTH eXCeeDINg TWO TIMeS bODY LeNgTH (TAL /BL 2.21���2.95, MeAN 2.60; N =4), WITH A POINTeD TAIL TIP. TAIL MUSCULATURe DISTINCT (MTH/BH 0.52���0.71, MeAN 0.62; N =4), RUNNINg PARALLeL fROM TAIL bASe TO ANTeRIOR TWO THIRDS Of TAIL LeNgTH, THeN gRADUALLY TAPeRINg TO THe TIP Of TAIL. TAIL fINS ReLATIVeLY LOW; UPPeR TAIL fIN STARTS POSTeRIORLY Of TAIL bASe, UPPeR TAIL fIN eDge SLIgHTLY CONVeX; bOTH UPPeR AND LOWeR TAIL fINS ReACH MAXIMUM HeIgHT IN POSTeRIOR THIRD. Oral disc. CUP-LIKe ORAL DISC ANTeROVeNTRALLY POSITIONeD (ODW/BW 0.32���0.37, MeAN 0.34; N =4), fRINgeD WITH SHORT CONICAL PAPILLAe; KeRATODONT ROW fORMULA: 4(2���4)/3(1���2), WITH AbOUT TWO THIN SPIKe-LIKe KeRATODONTS PeR 0.1 MM IN THe STAge 38 (FIg. 5E); jAW SHeATHS bLACK, RObUST; UPPeR jAW AND LOWeR jAW SHeATHS ReMARKAbLY DeVeLOPeD WITH DISTINCTLY SeRRATeD eDgeS (FIg. 5D). Coloration in life. BODY OLIVe-bROWN TO gReY WITH DISTINCT WHITISH-gOLDeN SPeCKLINg ON DORSAL AND LATeRAL SURfACeS. IRIS DARK, MARbLeD, WITH fOUR bLACK STRIPeS AT THe eXTeRNAL MARgIN. VeNTRAL SIDe Of bODY TRANSLUCeNT, WITH INTeSTINAL SPIRAL CLeARLY VISIbLe. INTeRNAL gILLS DISTINCTLY ReDDISH. TAIL fIN TRANSLUCeNT, gReYISH-bROWN (FIg. 5A���C). Coloration in preservative. WHeN PReSeRVeD IN 5% SOLUTION Of fORMALIN, COLORATION fADeS SIgNIfICANTLY, bODY TURNS SeMI-TRANSLUCeNT, DORSALLY AND DORSOLATeRALLY LIgHT bROWN, VeNTRALLY TRANSLUCeNT, TURNINg WHITISH ON THROAT AND CHeST RegION. TAIL MUSCULATURe fADeS TO gReY, UPPeR AND LOWeR TAIL fIN ALMOST TRANSLUCeNT WITH SLIgHT INDISTINCT gReY PIgMeNTATION. DORSOLATeRAL PARTS Of HINDLIMbS WITH WHITISH TO gReYISH PIgMeNTATION. Measurements. THe VARIATION Of MORPHOLOgICAL CHARACTeRS Of THe TADPOLeS IS SHOWN IN TAbLe 5. Comparisons. Leptolalax rowleyae sp. nov. bOTH MORPHOLOgICALLY AND MOLeCULARLY IS MOST SIMILAR TO THe MeMbeRS Of THe SMALL-bODIeD L. applebyi SPeCIeS gROUP INHAbITINg IN THe CeNTRAL HIgHLANDS Of CeNTRAL AND SOUTHeRN VIeTNAM AND THe NORTHeASTeRN PART Of CAMbODIA INCLUDINg L. applebyi, L. ardens, L. bidoupensis, L. kalonensis, L. maculosus, L. melicus, L. pallidus, L. pyrrhops, AND L. tadungensis. COMPARISONS Of THe NeW SPeCIeS WITH THe MeMbeRS Of L. applebyi SPeCIeS gROUP APPeAR TO be THe MOST PeRTINeNT. Leptolalax rowleyae sp. nov. CAN be DISTINgUISHeD fROM ALL OTHeR SPeCIeS Of Leptolalax (NON-MeMbeRS Of L. applebyi SPeCIeS gROUP) RePORTeD fOR INDOCHINA AND SURROUNDINg AReAS IN ITS OVeRALL MORPHOLOgY AS fOLLOWS: IN HAVINg SMALL bODY SIZe (SVL 23.4���25.4 MM IN ADULT MALeS, 27.0���27.8 MM IN ADULT feMALeS), Leptolalax rowleyae sp. nov. CAN be DISTINgUISHeD fROM THe LARgeR CONgeNeRS INCLUDINg L. bourreti (MALeS 28.0���36.2 MM, feMALeS 42.0��� 45.0 MM), L. botsfordi (MALeS 29.1���32.6 MM, feMALeS 30.0���31.8 MM), L. eos (MALeS 33.1���34.7 MM, feMALe 40.7 MM), L. firthi (MALeS 26.4���29.2 MM, feMALeS 25.7���36.9 MM), L. fuliginosus (MALeS 28.2���30.0 MM), L. minimus (MALeS 25.7���31.4 MM, feMALeS 31.6���37.3 MM), L. nahangensis (MALe 40.8 MM), L. nyx (MALeS 26���30 MM, feMALeS 36���42 MM), L. pelodytoides (MALeS 27.5���32.3 MM, feMALeS 35.5���37.8 MM), L. petrops (MALeS 23.6���27.6 MM, feMALeS 30.3���47.0 MM); L. platycephalus (MALe 35.1 MM, feMALe 46.0 MM), L. puhoatensis (MALeS 24.2���28.1 MM, feMALeS 27.3���31.5 MM), L. solus (MALe 27.6 MM), AND L. sungi (MALeS 48.3���52.7 MM, feMALeS 56.7���58.9 MM), AND fROM THe SMALLeR SPeCIeS INCLUDINg L. kecil (MALeS 19.3���20.5 MM, feMALe 25 MM) AND L. pluvialis (MALeS 21.3���22.3 MM). IN HAVINg PINKISH MILK-WHITe TO LIgHTLY bROWN CHeST AND beLLY WITH NUMeROUS WHITe SPeCKLeS, THe NeW SPeCIeS ALSO DIffeRS fROM L. aereus, L. bourreti, L. firthi, L. fuliginosus, L. mininus, L. nahangensis, L. nyx, L. pelodytoides, L. solus, L. sungi, L. tuberosus, AND L. zhangyapingi (ALL Of WHICH HAVe MOSTLY WHITe, CReAMY WHITe OR PALe gReY VeNTeRS, WITHOUT WHITe SPeCKLINg AND WITH OR WITHOUT DARK SPOTS OR MOTTLINg); fROM L. botsfordi (VS. ReDDISH-bROWN VeNTRAL SURfACeS WITH WHITe SPeCKLINg); fROM L. croceus (VS. ORANge beLLY); fROM L. eos (VS. CReAMY WHITe beLLY WITH bROWN SPeCKLINg ALONg ITS MARgINS); fROM L. petrops (VS. PALe PINK AND SLIgHTLY TRANSLUCeNT beLLY, PARTICULARLY AT eITHeR SIDe Of THROAT; eDgeS Of THROAT PALe bROWN WITH WHITe SPeCKLINg, CONCeNTRATeD TOWARDS SNOUT; VeNTRAL SURfACe Of CHeST AND AbDOMeN IMMACULATe WHITe); fROM L. pluvialis (VS. gReY VeNTeR WITH DARK gReY MARbLINg, UNIfORM PALe gReY THROAT WITH SPeCKLINg AROUND ITS MARgINS); fROM L. puhoatensis (VS. ReDDISH-bROWN, OfTeN WITH fAINT WHITe DUSTINg IN MALeS AND A PALe PINK IN feMALeS); fROM L. melanoleucus AND L. ventripunctatus (bOTH DISPLAY LARge PATCHeS Of DISTINCT bROWN AND WHITe MARbLINg); fROM L. heteropus (VS. gReY VeNTeR, SPeCKLeD WITH bLACK); AND fROM L. kecil (VS. A UNIfORMLY DARK VeNTeR WITH LARge, DARK ORANge PeCTORAL gLANDS). BY HAVINg bLACK MARKINgS (SPOTS OR bLOTCHeS) ON THe fLANKS, THe NeW SPeCIeS CAN be fURTHeR DIffeReNTIATeD fROM L. aereus, L. croceus, L. eos, L. firthi, L. isos, AND L. tuberosus, ALL Of WHICH DO NOT HAVe THe MARKINgS. IN HAVINg A DISTINCT TYMPANUM, THe NeW SPeCIeS CAN be eASILY DIAgNOSeD fROM L. croceus AND L. tuberosus (THeSe TWO SPeCIeS TYMPANUM IS HIDDeN) AND fROM L. sungi (WHICH IS RePORTeD TO LACK eXTeRNAL TYMPANUM, HOWeVeR IN SOMe POPULATIONS AN INDISTINCT TYMPANUM IS STILL DISCeRNIbLe). IN HAVINg TOeS WITH bASAL WebbINg AND NO LATeRAL fRINgINg, Leptolalax rowleyae sp. nov. CAN be DISTINgUISHAbLe fROM L. eos, L. firthi, AND L. isos, ALL Of WHICH HAVe MORe eXTeNSIVe TOe WebbINg AND DISTINCT LATeRAL fRINgeS ON TOeS. IN HAVINg DORSAL SKIN MOSTLY SMOOTH WITH NUMeROUS TINY TUbeRCLeS, THe NeW SPeCIeS IS ALSO DISTINgUISHAbLe fROM L. croceus, L. isos, L. minimus, L. petrops, L. solus, L. tuberosus, AND L. ventripunctatus (ALL THeSe SPeCIeS HAVe ROUgHLY gRANULAR OR TUbeRCUL, Published as part of Nguyen, Luan Thanh, Dzung Trung Le, Nikolay A. Poyarkov Jr., Vo, Ba Dinh, Phan, Hoa Thi, Duong, Tang Van, Murphy, Robert W. & Nguyen, Sang Ngoc, 2018, A new species of Leptolalax (Anura: Megophryidae) from Son Tra Peninsula, central Vietnam, pp. 1-21 in Zootaxa 4388 (1) on pages 7-17, DOI: 10.11646/zootaxa.4388.1.1, http://zenodo.org/record/1187777, {"references":["Gosner, K. L. (1960) A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16, 183 - 190."]}

Oligodon culaochamensis sp. nov. (Figs. 2���5) Holotype. ITBCZ 5646, adult male, collected from Hon Lao, Cu Lao Cham Islands, Quang Nam Province, Vietnam; coordinates 15��56���38���N, 108��30���44���E; elevation 41 m a.s.l. by L.T. Nguyen, V.D.H. Nguyen, and S.N. Nguyen on 27 July 2016. Paratypes. Five specimens, also collected from Hon Lao, Cu Lao Cham Islands. ITBCZ 5618, adult females, collected on 21 July 2016, coordinates 15��56���27���N, 108��31���13���E; elevation 106 m a.s.l.; ITBCZ 5631, adult male, collected on 24 June 2016, coordinates 15��56���40���N, 108��30���44���E; elevation 68 m a.s.l. by L.T. Nguyen, V.D.H. Nguyen, and S.N. Nguyen; ITBCZ 5655���6, two adult females, and ITBCZ 5658, a juvenile, collected in 2012 by L.T. Nguyen and H.T. Phan. (Figs. 4 & 5) Diagnosis. Oligodon culaochamensis sp. nov. is distinguished from its congeners by a combination of the following morphological characters: (1) medium size in adults (TL �� 582 mm); (2) 17 or 19 dorsal scale rows at neck and midbody and 15 or 17 rows before vent; (3) LaL/TL ratio 0.218���0.219 in males, 0.166���0.169 in females; (4) ventrals 167���169 in males and 179���182 in females; (5) subcaudals 63���66 in males and 51���52 in females; (6) presubocular present; (7) 8 supralabials, fourth and fifth entering orbit; (9) 9���10 maxillary teeth, posterior three enlarged; (10) nasal divided; (11) temporal streak absent; (12) long and deeply forked hemipenes, extending to 20th subcaudal, without spines and papillae but with a prominent diagonal ridge, oblique flounces and distal calyces; (13) cloacal plate undivided; (14) 10���11+3���4 dorsal blotches; and (15) vertebral stripe on tail and anterior part of body. Description of holotype. Adult male; head small and elongate (HW/HL = 0.579), faintly distinct from neck; body rather slender; SVL 424 mm; tail quite long (TaL/TL = 0.219), 119 mm in length; eye small with round pupil, eye diameter equal to distance between eye and nostril; snout quite pointed with a large oval nostril piercing the upper part of nasal. Head scalation. Rostral thick and curved on to upper snout surface, visible from above, pointed posteriorly, touching nasals, internasals and first supralabial on both sides; the portion of rostral visible from above shorter than its distance from frontal; supralabials 8/8, the 4th and 5th bordering eye; 1/1 loreal; 1/1 presubocular; 1/1 preocular; 2/2 postoculars; nasal divided vertically; 2+2/2+2 temporals, anterior ones elongated; prefrontal wider than long and larger than internasal; frontal large and hexagonal, longer than wide; parietals larger than frontals, bordered laterally by the upper temporals and anteriorly by frontal, supraoculars and postoculars; 9/9 infralabials, first pair medially in contact with each other, first four in contact with anterior chin shield, the 5th largest and touching posterior chin shield; two pairs of gular scales between posterior chin shield and first ventral. Body scalation. Dorsal scales smooth, in 17���17���15 rows; scale row reductions from 17 to 16 at ventral 135 and from 16 to 15 at ventral 141; vertebral scales similar to other dorsal scales in size and shape; ventrals 167; cloacal plate entire; subcaudals 63, all paired; terminal caudal scale forming a pointed cap. Dentition. Maxillary teeth 9, curved posteriorly, smaller and shorter anteriorly; posterior three being enlarged (Fig. 2); palatine and pterygoid teeth developed and slightly curved posteriorly. Hemipenis. Hemipenis long with two twisted lobes, deeply forked at the level of 6th SC and tip of each organ reaching the SC 20 (Fig. 3), without spines and papillae; each lobe with numerous smooth, small and oblique folds, which a deep diagonal groove interrupts; folds form smooth hexagonal calyces at the distal part of the organ (Fig. 3); prominent diagonal ridge starting from in front of the forked point and extending to near the tip; diagonal ridge easily visible when pressing the tail base to evert the hemipenis (Fig. 3, top); surface of the ridge with calyces changes directly from the oblique folds; hemipenes becoming twisted in everted condition when water injected inside. Coloration. In life, overall dorsal coloration brown to pinkish brown with 10 darker blotches on body and 3 on tail. Body blotches about 3���4 dorsal scales wide. Numerous scattered dorsal black spots confined to edges of the scales. Edge of each scale and intervening skin pink. Bright vertebral strip on anterior part of body and tail. A dark brown band from eye to eye through preoculars, supraoculars, frontal, prefrontals and internasals, continues over the 4th and 5th supralabials ending at lower part of the 6th supralabial. Head with another dark brown arrow-shaped marking, apex pointing forward and reaching the posterior part of frontal, backwards obliquely crossing neck and nearly reaching the 6th and 7th ventrals. Temporal streak absent. Snout, lateral and lower parts of head white. Ventral color cream to pink or reddish with black rectangular blotches. Ventral surface of tail white with black spots. Iris black and round; remaining part of the eye somewhat bicolor diagonally; anterior part with uniform color streak, posterior part mainly bright yellow. In alcohol, color becoming paler, but the pattern remains; iris white; venter cream to white with black blotches and dots, pink or reddish color disappears. Variation. Table 1 summarizes variation in size and scalation of the holotype and five paratypes. In life, paratype ITBCZ 5618 has a pink edge on each dorsal scale and the intervening skin, as occurs in the holotype. In comparison, the edge of each dorsal scale and the intervening skin of paratype ITBCZ 5631 is white and its dorsal coloration is grey. Color in life of the other paratypes was not recorded. ITBCZ 5618 has a vertebral stripe not only on tail and anterior part of body but also on posterior part of body (Fig. 5 A). The juvenile paratype (ITBCZ 5658, Fig. 5 D) is similar to the adults in coloration and patterned. The ventral surface of the tails of two paratypes (ITBCZ 5618 and ITBCZ 5656) are white without grey spots. The most variable character is the number of dorsal scale rows. The paratype (ITBCZ 5655) has 17���19���17 dorsal scale rows, increasing from 17 to 18 at VS 16 and to 19 at VS 33, reducing from 19 to 18 at VS 102 and to 17 at VS 106. The paratype (ITBCZ 5656) has 19���19���17 dorsal scale rows, the first and second reductions occur at VS 105 and 110, respectively. The paratype (ITBCZ 5631) has 18���17���15 dorsal scale rows, reducing from 18 to 17 at VS 18, increasing from 17 to 18 at VS 40, from 18 to 19 at VS 42, reducing from 19 to 18 at VS 78, from 18 to 17 at VS 82, from 17 to 16 at VS 153 and from 16 to 15 at 158, and increasing from 15 to 16 near the vent at VS 162. The paratype (ITBCZ 5618) has 17���19���17 dorsal scale rows, increasing from 17 to 18 at VS 22 and to 19 at VS 24, reducing from 19 to 18 at VS 103 and to 17 at VS 105. Hemipenis of the second male (paratype ITBCZ 5631, Fig. 4) is similar to that of the holotype. It also extends to SC 20 and twisted, but deeply forked at the level of 7th SC. Natural history. All specimens were collected at night and on the ground in dry primary forests or plantations, usually near the bank of small streams. The stomach of the holotype contained two soft shell eggs (size 15 mm x 7 mm). These eggs probably belong to a snake or lizard. Sexual dimorphism. Males have longer tails than females (male TaL/TL = 0.218���0.219 vs. 0.166���0.169), a larger size (maximal SVL = 544 mm vs. 383 mm), more subcaudals (SC 63���66 vs. 51���52) but fewer ventrals (VS 167���169 vs. 179���182). Distribution. The new species is currently known only from Cu Lao Cham Islands, Quang Nam Province, Vietnam. Etymology. The specific epithet culaochamensis is derived from Cu Lao Cham Islands, where the new species was discovered. Comparisons. Based on hemipenal morphology (deeply forked, no papillae, no spines), O. culaochamensis sp. nov. is a member of the O. cyclurus group as defined and mentioned in previous publication (Smith 1943, David et al. 2008a, 2008b; Green et al. 2010; Neang et al. 2012; Nguyen et al. 2016). Therefore, the new species differs from all congeners within the Oligodon taeniatus group (David et al. 2008b; Neang et al. 2012; Nguyen et al. 2016), which includes O. barroni (Smith), O. deuvei David, Vogel & Rooijen, O. moricei David, Vogel & Rooijen, O. mouhoti (Boulenger), O. pseudotaeniatus David, Vogel & Rooijen, and O. taeniatus (G��nther), because they have obvious papillae on hemipenes, except for O. moricei in which males remain unknown; from all congeners within the O. cinereus group (David et al. 2008a, 2012; Green et al. 2010; Neang et al. 2012; Vassilieva 2015; Nguyen et al. 2016), which includes O. albocinctus (Cantor), O. cinereus (G��nther), O. inornatus (Boulenger), O. joynsoni (Smith), O. maculatus (Taylor), O. melanozonatus Wall, O. nagao David, Nguyen, Nguyen, Jiang, Chen, Teynie & Ziegler, O. splendidus (G��nther) and O. woodmasoni (Sclater), because they do not have forked hemipenis, except for O. melanozonatus, in which the hemipenis remains unknown; from all congeners within the O. dorsalis group (Smith 1943, Vassilieva 2015; Nguyen et al. 2016), which includes O. catenatus (Blyth), O. dorsalis (Gray & Hardwicke), O. eberhardti Pellegrin, O. erythrogaster Boulenger, O. hamptoni Boulenger, O. lacroixi Angel & Bourret, and O. mcdougalli Wall, because they have unforked or slightly forked hemipenis, except for O. lacroixi, in which the hemipenis is unknown; from all congeners within the O. torquatus group (Green et al. 2010, Smith 1943, Vassilieva 2015; Nguyen et al. 2016), which includes O. cruentatus (G��nther), O. planiceps (Boulenger), O. theobaldi (G��nther), and O. torquatus (Boulenger), because they have unforked hemipenis with papillae. The new species can be distinguished from O. moricei, which is known from a female only, by having dorsal blotches (vs. absent), longer tail (TaL/TL = 0.166���0.169 vs 0.113), more subcaudals (51���52 vs 41), fewer maxillary teeth (9���10 vs. 12) and a presubocular (vs absent); from O. melanozonatus by having dorsal blotches (vs. cross-bars), a loreal (vs. absent), a single cloacal plate (vs. divided), more supralabials (8 vs. 6), and more subcaudals (51���66 vs. 42���45); and from O. lacroixi by having an entire cloacal plate (vs. divided), more scale rows at midbody (17���19 vs. 15), more supralabials (8 vs. 5) and a loreal (vs. absent). Oligodon culaochamensis sp. nov. differs from other congeners in the Indochinese region (Vietnam, Laos, Cambodia, Thailand, Myanmar, Peninsular Malaysia, and southern China [Yunnan]), except for those of the O. cyclurus group, as follows: from O. annamensis Leviton by having more dorsal scale rows (17���19 at neck and midbody and 15 before vent vs. 13���13���13), more supralabials (8 vs. 6), more maxillary teeth (9���10 vs. 8) and by dorsal color pattern (blotches vs. cross-bars); from O. arenarius by having deeply forked hemipenis (vs. unforked), a larger size (582 mm vs. 389 mm), more maxillary teeth (9���10 vs. 6���8) and more ventrals (167���183 vs. 131���144); from O. booliati Leong & Grismer by having more supralabials (8 vs. 6 or 7), more ventrals (167���182 vs. 143���153) and dorsal color pattern (blotches vs. cross-bars); from O. jintakunei Pauwels, Wallach, David & Chanhome by having more dorsal scale rows at midbody (17���19 vs. 15), fewer ventrals (167���182 vs. 189), entire cloacal plate (vs. divided) and dorsal pattern (blotches vs. cross-bars) (Pauwels et al. 2002; Vassilieva 2015); from O. octolineatus (Schneider) by having a smaller size (582 mm vs. 780 mm), more supralabials (8 vs. 6), the 4th and 5th supralabials entering the eye (vs. SL 3rd and 4th) and dorsal color pattern (blotches vs. broad stripes); from O. purpurascens (Schlegel) by having a smaller size (TL 582 mm vs. 790 mm), portion of rostral visible from above much shorter (vs. as long as) than its distance from the frontal, dorsal color pattern (blotches vs. transverse bands or cross-bars) and the absence of temporal streak (vs. presence); and from O. vertebralis G��nther by having more supralabials (8 vs. 7), SL 4th and 5th entering the eye (vs. SL 3rd and 4th), more dorsal scale rows at midbody (17���19 vs. 15), and an entire cloacal plate (vs. divided). Oligodon culaochamensis sp. nov. differs from all other congeners within the O. cyclurus group by the combination of the following characters: maximum size, TaL/TL ratio, ventrals, subcaudals, maxillary teeth, length and shape of hemipenis, dorsal pattern and temporal streak (see Table 2). The new species is different from O. condaoensis by having a longer tail in males (TaL/TL = 0.218���0.219 vs. 0.129���0.132) and females (0.166���0.169 vs 0.126), more subcaudals in males (63���66 vs. 37) and females (51���52 vs. 33���34), more anterior temporal (2 vs. 1), a presubocular (vs. absent), fewer maxillary teeth (9���10 vs. 11���13), different hemipenis (LHSC = 20 vs. 13���14, presence of oblique folds and diagonal ridge vs. absent) and different dorsal color pattern (blotches vs. uniform or faint tripe); from O. cattienensis by having a longer tail in males (TaL/ TL = 0.218���0.219 vs. 0.110���0.130) and females (0.166���0.169 vs 0.110), more ventrals in females (179���182 vs. 168), more subcaudals in males (63���66 vs. 32���36) and females (51���52 vs. 32), fewer maxillary teeth (9���10 vs. 11��� 12), fewer number of dorsal blotches (10���11+3���4 vs. 24���33+5), different head pattern (absence [vs. presence] of temporal streak) and different hemipenis shape (deeply forked vs. forked near the tip); from O. cyclurus (Cantor) by having a smaller size (582 mm vs. 710 mm), longer tail in males (TaL/TL = 0.218���0.219 vs. 0.127���0.156) and females (0.166���0.169 vs 0.107���0.138), more ventrals in females (179���182 vs. 168���172), more subcaudals in males (63���66 vs. 37���43) and females (51���52 vs. 30���44), a longer hemipenis (LHSC = 20 vs. 15���18), different dorsal pattern (presence [vs. absence] of vertebral stripe on tail and anterior part of body) and different head pattern 2008a, Neang et al. 2012, Nguyen et al. 2016, Pope 1935, Smith 1943, Vassilieva et al. 2013, Wall 1923 and examined specimens. See Material and Methods for abbreviations; *, values given species, not for males or females. (absence [vs. presence] of temporal streak); from O. fasciolatus (G��nther) by having a smaller size (582 mm vs. 807 mm), fewer dorsal scale rows at neck (17���19 vs. 21���23) and midbody (17���19 vs. 21), more subcaudals in males (63���66 vs. 43���61) and females (51���52 vs. 34���48), different dorsal pattern (presence [vs. absence] of vertebral stripe on tail and anterior part of body) and different head pattern (absence [vs. presence] of temporal streak); from O. formosanus (G��nther) by having a smaller size (582 mm vs. 942 mm), longer tail in males (TaL/ TL = 0.218���0.219 vs. 0.165���0.195), more subcaudals in males (63���66 vs. 49���55) and females (51���52 vs. 43���48), different hemipenis length (LHSC = 20 vs. 15���18) and shape (short papillae absent [vs. present]), different dorsal pattern (blotches vs. reticulate) and different head pattern (absence [vs. presence] of temporal streak); from O. juglandifer (Wall) by having a smaller size (582 mm vs. 853 mm), more supralabials (8 vs. 7) and the SL 4th and 5th entering the eye (vs. SL 3rd and 4th), different dorsal pattern (blotches vs. bands, presence [vs. absence] of dorsal stripe) and different head pattern (absence [vs. presence] of temporal streak); from O. kampucheaensis Neang, Grismer & Dattry by having a longer tail in males (TaL/TL = 0.218���0.219 vs. 0.151), more dorsal scale rows at neck and midbody (17���19 vs. 15), more ventrals (167���169 vs. 165) and subcaudals (63���66 vs. 39) in males, a longer hemipenis (LHSC = 20 vs. 11), different dorsal pattern (blotches vs. cross-bars, presence [vs. absence] of dorsal stripe) and different head pattern (absence [vs. presence] of temporal streak); from O. kheriensis Acharji & Ray by having a longer tail (TaL/TL = 0.166���0.219 vs. 0.121���0.161), fewer ventrals (167���182 vs. 196), more subcaudals (51���66 vs. 38���43), and different dorsal pattern (blotches vs. uniform and presence [vs. absence] of dorsal stripe); from O. macrurus (Angel) by having a shorter tail (TaL/TL = 0.218���0.219 vs. 0.240), more ventrals (167���182 vs. 143���152), fewer subcaudals (51���66 vs. 76���83), a shorter hemipenis (LHSC = 20 vs. 29), dorsal pattern (blotches vs. uniform and presence [vs. absence] of dorsal stripe) and head pattern (absence [vs. presence] of temporal streak); from O. ocellatus (Morice) by having a smaller size (582 mm vs. 825 mm), a remarkably longer tail in both males (TaL/TL = 0.218���0.219 vs. 0.112���0.141) and females (TaL/TL = 0.166���0.169 vs. 0.094��� 0.114), more ventrals in males (167���169 vs. 156���165), more subcaudals in both males (63���66 vs. 32���44) and females (51���52 vs. 26���33), a longer hemipenis (LHSC = 20 vs. 15���17), dorsal pattern (presence [vs. absence] of dorsal stripe), and head pattern (absence [vs. presence] of temporal streak); and from O. saintgironsi by having a smaller size (582 mm vs. 676 mm), fewer ventrals in females (179���182 vs. 184), more subcaudals in males (63���66 vs. 55���59), a shorter hemipenis (LHSC = 20 vs. 27���28), hemipenis shape (oblique folds or flounces vs. large calyces throughout; presence [vs. absence] of diagonal ridge) and head pattern (absence [vs. presence] of temporal streak). Oligodon culaochamensis sp. nov. is similar to O. chinensis (G��nther) in having hemipenis with oblique folds and a prominent diagonal ridge (Pope 1935; Smith 1943). However, the new species can be distinguished from O. chinensis by having a smaller size (582 mm vs. 729 mm), longer tail in males (TaL/TL = 0.218���0.219 vs. 0.187��� 0.195), fewer ventrals in both males (167���169 vs. 175���184) and females (179���182 vs. 182���206), presubocular (vs. usually no presubocular), two anterior temporals (vs. usually one), remarkably longer hemipenis (LHSC = 20 vs. 12���14), dorsal pattern (blotches vs. cross-bars) and head pattern (absence [vs. presence] of interrupted temporal streak)., Published as part of Nguyen, Sang Ngoc, Nguyen, Luan Thanh, Nguyen, Vu Dang Hoang, Phan, Hoa Thi, Jiang, Ke & Murphy, Robert W., 2017, A new species of the genus Oligodon Fitzinger, 1826 (Squamata: Colubridae) from Cu Lao Cham Islands, central Vietnam, pp. 333-346 in Zootaxa 4286 (3) on pages 335-343, DOI: 10.11646/zootaxa.4286.3.2, http://zenodo.org/record/828537, {"references":["Smith, M. A. (1943) The Fauna of British India, Ceylon and Burma, including the whole of the Indo-Chinese subregion. Reptiles and Amphibians, VOl. III-Serpentes. TaylOr and Francis Ltd, LOndOn, 525 pp.","David, P., VOgel, G. & Pauwels, O. S. G. (2008 a) A new species Of the genus Oligodon Fitzinger, 1826 (Squamata: COlubridae) frOm sOuthern Vietnam and CambOdia. Zootaxa, 1939, 19 - 37.","David, P., VOgel, G. & van ROOijen, J. (2008 b) A revisiOn Of the Oligodon taeniatus (Gunther, 1861) grOup (Squamata: COlubridae), with the descriptiOn Of three new species frOm the IndOchinese regiOn. Zootaxa, 1965, 1 - 49.","Green, M. D., OrlOv, N. L. & Murphy, R. w. (2010) TOward a phylOgeny Of the kukri snakes, genus Oligodon. Asian Herpetological Research, 1, 1 - 21.","Neang, T., Grismer, L. L. & Daltry, J. C. (2012) A new species Of kukri snake (COlubridae: Oligodon Fitzinger, 1826) frOm the PhnOm SamkOs wildlife Sanctuary, CardamOm MOuntains, sOuthwest CambOdia. Zootaxa, 3388, 41 - 55.","Nguyen, S. N., Nguyen, V. D. H., Le, S. H. & Murphy, R. w. (2016) A new species Of kukri snake (Squamata: COlubridae: Oligodon Fitzinger, 1826) frOm COn DaO Islands, sOuthern Vietnam. Zootaxa, 4139 (2), 261 - 273. https: // dOi. Org / 10.11646 / zOOtaxa. 4139.2.9","David, P., Nguyen, T. Q., Nguyen, T. T., Jiang, K., Chen, T., Teynie, A. & Ziegler, T. (2012) A new species Of the genus Oligodon Fitzinger, 1826 (Squamata: COlubridae) frOm nOrthern Vietnam, sOuthern China and central LaOs. Zootaxa, 3498, 45 - 62.","Vassilieva, A. B. (2015) A new species Of the genus Oligodon Fitzinger, 1826 (Squamata: COlubridae) frOm cOastal sOuthern Vietnam. Zootaxa, 4058 (2), 211 - 226.","Pauwels, O. S. G., wallach, V., David, P. & ChanhOme, L. (2002) A new species Of Oligodon Fitzinger, 1826 (Serpentes, COlubridae) frOm sOuthern Peninsular Thailand. The Natural History Journal of Chulalongkorn University, 2, 7 - 18.","POpe, C. H. (1935) The Reptiles of China. The American Museum Of Natural HistOry, New YOrk, xlii + 604 pp., 27 pls.","Vassilieva, A. B., Geissler, P., GalOyan, E. A., POyarkOv, N. A., van Devender, R. w. & Bohme, w. (2013) A new species Of kukri snake (Oligodon Fitzinger, 1826; Squamata: COlubridae) frOm the Cat Tien NatiOnal Park, sOuthern Vietnam. Zootaxa, 3702 (3), 233 - 246."]}

Oligodon condaoensis sp. nov. (Figs. 2 & 3) Holotype. ITBCZ 2595, adult male, collected from Hon Ba Island, Con Dao District, Ba Ria���Vung Tau Prov., Vietnam; coordinates 8��39���03���N, 106��33���29���E; elevation 15 m a.s.l. by Sang Ngoc Nguyen on 24 October 2015. Paratypes. ITBCZ 2596, adult male; ITBCZ 2597���8, two adult females; collected around the site of the holotype on Hon Ba Island by Sang Ngoc Nguyen on 24 and 25 October 2015. Diagnosis. Oligodon condaoensis sp. nov. is distinguished from all of its congeners by a combination of the following morphological characters: (1) medium size in adults (total length up to 552 mm); (2) dorsal scale rows 17���17���15; (3) deeply forked hemipenes without papillae and spines, extending to SC 13 or 14; (4) maxillary teeth 11���13, the posterior three being enlarged; (5) cloacal plate undivided; (6) head scalation complete; (7) nasal divided; (8) presubocular absent; (9) ventrals 168���176; (10) subcaudals 33���37; (11) overall dorsal coloration dark gray, faint body stripes present or absent; (12) venter cream to dark gray without rectangular blotches. Description of holotype. Adult male; body robust, rather elongate; SVL 461 mm; tail quite short (TL/SVL = 0.15) and robust, 70 mm in length; head short (HL/SVL = 0.04), faintly distinct from neck; eye small (ED/HH = 0.35) with round pupil, eye diameter equal to distance between eye and nostril; snout obtuse; large oval nostril piercing the upper part of nasal. Head scalation. Rostral high 3.37 mm, wide 3.57 mm, thick and curved on to upper snout surface, well visible from above, pointed posteriorly, touching internasals, nasals, and first supralabial on each side; supralabials 8/8, the 4th and 5th bordering eye, the 7th being largest; 1/1 loreal; presubocular absent; nasal divided vertically; 1/1 preocular; 2/2 postoculars; 1+2/1+2 temporals, anterior one elongated; prefrontal large, somewhat hexagonal, wider than long; frontal large and pentagonal, slightly longer than wide; parietals larger than frontals, bordered laterally by the first and the upper second temporals and posteriorly by four small scales; 8/8 infralabials, fist pair medially in contact with each other, first four InLs in contact with anterior chin shield, the 5th largest and touching posterior chin shield. Body scalation. Dorsal scales smooth, in 17���17���15 rows; scale row reductions from 17 to 15 around the position corresponding to ventral 98th; vertebral scales similar to other dorsal scales in size and shape; ventrals 172; cloacal plate entire; subcaudals 37, all paired; terminal caudal scale forming a pointed cap. Dentition. Maxillary teeth 13, curved posteriorly, the posterior three being enlarged (Fig. 2 C); palatine and pterygoid teeth 7 and 14, respectively, slightly curved posteriorly (Fig. 2 D). Hemipenis. Hemipenis with two enlarged lobes, deeply forked at the level of 6th SC and tip of each organ reaching the SC 13th; proximal part with faint folds forming more or less hexagonal cells, these cells fainter at distal part (Fig. 2 E���G). Coloration. In life, overall dorsal coloration dark gray, including dorsal, lateral scales, and lateral sides of ventral scales. No distinct stripes or blotches. Head more or less uniform dark gray. Infralabials, lower part of supralabials, and rostral whitish. Underside of head and anterior part of body white to cream. The ventral color of body changes backwardly from cream anteriorly to dark gray dots or blotches and then to dark gray posteriorly, not forming rectangular blotches. Two white stripes along ventral-lateral part of ventral scales. Ventral surface of tail white with some gray dots. Pupil round and black. In alcohol, the pattern paler, becoming gray. Pupil white. Variation. Table 1 summarizes variation in size and scalation. Head scalation complete but presubocular absent in all specimens. In the three paratypes, brighter longitudinal stripes are faint in life and in alcohol. A narrow grayish vertebral stripe extends from neck to the vent level, the width of this stripe is narrower than the width of the vertebral scale but in the neck the stripe width expanding to about two dorsal scales; paravertebral stripes very faint; a faint but broad, grayish stripe extends up to the vent on dorsal scale rows 4���6. In general, the faint stripe pattern of the new species, if present, is as follows: lateral side of ventral scale and DS 1���3 dark gray or gray; DS 4���6 brighter; DS 7���8 dark gray or gray; and vertebral scale brighter (Fig. 3 C). Paratypes ITBCZ 2597���8 with a dark oblique streak below the eye, on SL 5���6; frontal and parietals with faint blotches. Ventral pattern of paratype ITBCZ 2596 cream to white with few gray dots; paratypes ITBCZ 2597���8 similar to that of the holotype. Hemipenis of the paratype ITBCZ 2596 similar to that of the holotype. The palatine and pterygoid teeth of the paratypes ITBCZ 2596���8 are 6+12, 7+13, and 7+12, respectively. The posterior three maxillary teeth of the paratypes are also enlarged. Natural history. All specimens were collected in the daytime, between 9:00 am and 12:00 pm. The animals were found active on the ground of evergreen forest (Fig. 3 D) near by Hon Ba Forest Ranger Station. Two nights of searching in the area failed to find the snake. This infers that the new species is diurnal. Another species, O. fasciolatus, was recorded sympatrically with the new species. Sexual dimorphism. The males are larger (SVL 461���481 mm vs. 395���401 mm) and darker than the females. The faint stripes on females are more visible than on the males. Tails in males are longer than in females (SC 37 vs. 33���34; TL/SVL 0.15 vs. 0.14). Distribution. The new species is currently known only from Hon Ba Island, Con Dao District, Ba Ria���Vung Tau Prov., Vietnam. Etymology. The specific epithet condaoensis is derived from Con Dao Islands, where the new species was discovered. Comparisons. Oligodon condaoensis sp. nov. differs from all other congeners within the Oligodon taeniatus group, the O. cinereus group, the O. cyclurus group (David et al. 2008 a, b; Green et al. 2010; Neang et al. 2012), the O. dorsalis group (Smith 1943, Vassilieva 2015), and the O. torquatus group (Green et al. 2010, Smith 1943, Vassilieva 2015), and other species of the genus in Indochina by the combination of the following characters: body and head scalation, maxillary teeth, hemipenis, body size, and dorsal color pattern. By having deeply forked hemipenis without obvious papillae, O. condaoensis sp. nov. differs from the O. taeniatus group (David et al. 2008b; Neang et al. 2012), including O. barroni (Smith), O. deuvei David, Vogel & Rooijen, O. moricei David, Vogel & Rooijen, O. mouhoti (Boulenger), O. pseudotaeniatus David, Vogel & Rooijen, and O. taeniatus (G��nther), except for O. moricei in which males remain unknown (Table 2). The new species is also distinguishable from O. barroni by having a larger size (TL 552 mm vs. 401 mm), a higher number of ventrals (168���176 vs. 136���160), and different dorsal color pattern (gray or faint stripes vs. butterfly-like blotches on dorsum and tail); from O. deuvei by having a higher number of ventral scales (168���176 vs. 140���155), different body and head color pattern (uniform or faint vertebral stripe vs. broad and bright vertebral stripe, uniform or faint blotches vs. four or five major markings, respectively); from O. moricei by having a lower number of subcaudals (33���34 in females vs. 41), a lower number of infralabials (8 vs. 9), a higher number of maxillary teeth (13 in females vs. 12), and different dorsal color pattern (uniform or faint dorsal stripes vs. broad vertebral stripe, clear paravertebral and dorsolateral stripes); from O. mouhoti by having a bigger size (TL 552 mm vs. 339 mm), a higher number of ventrals (168���176 vs. 145���163), lower number of infralabials (8 vs. 9 or 10) and maxillary teeth (11���13 vs. 14���16), a shorter hemipenis (extending to SC 13���14 vs. 18���19), and the absence of blotches on tail and clear marks on head (vs. presence); from O. pseudotaeniatus by having a bigger size (TL 552 mm vs. 320 mm), a higher number of ventrals (168���176 vs. 137���156), fewer maxillary teeth (11���13 vs. 15), and the absence of presubocular (vs. presence); from O. taeniatus by having a bigger size (TL 552 mm vs. 447 mm), fewer dorsal scale rows at neck and midbody (17���17���15 vs. 19���19���15), more ventrals (168���176 vs. 142���165), fewer infralabials (8 vs. 9), and the absence or faintness of body stripes (vs. presence). continued. Oligodon condaoensis sp. nov. differs from members of the O. cinereus group (David et al. 2008a, 2012; Green et al. 2010; Neang et al. 2012; Vassilieva 2015), including O. albocinctus (Cantor), O. cinereus (G��nther), O. inornatus (Boulenger), O. joynsoni (Smith), O. maculatus (Taylor), O. melanozonatus Wall, O. nagao David, Nguyen, Nguyen, Jiang, Chen, Teynie & Ziegler, O. splendidus (G��nther), and O. woodmasoni (Sclater), by having a deeply forked hemipenis (vs. unforked) except for O. melanozonatus, in which the hemipenis is unknown. The new species also differs from O. albocinctus by dorsal scale rows (17���17���15 vs. 19���17[19]���15[17]), fewer ventrals and subcaudals (168���176 vs. 177���208, 33���37 vs. 40���69, respectively), more supralabials (8 vs. 7), a shorter hemipenis (extending to SC 13���14 vs. 24), dark uniform or faint body stripes (vs. cross-bars); from O. cinereus by having smaller size (TL 552 mm vs. 700 mm) and dorsal color pattern (dark uniform or stripes vs. cross-bars [based on O. cinereus from Con Dao Islands, as also mentioned by Smith 1943: 216���217 as form IV and Campden- Main 1970]); from O. inornatus by dorsal scale rows (17���17���15 vs. 15���15���15); from O. joynsoni by having fewer ventrals and subcaudals (168���176 vs. 186���197, 33���37 vs. 43���50, respectively) and dorsal color pattern (dark uniform or faint stripes vs. blotches and reticulated cross-bars); from O. maculatus by having more ventrals (168���176 vs. 156���164) and different dorsal color pattern (dark uniform or strips vs. dark blotches); from O. melanozonatus by having loreals (vs. absence), an entire cloacal plate (vs. divided), fewer subcaudals (33���37 vs. 42���45), higher numbers of supralabials, infralabials, and maxillary teeth (8, 8, and 11���13 vs. 6, 6, and 8, respectively); from O. nagao by having a smaller size (TL 552 mm vs. 786 mm), fewer ventrals (168���176 vs. 184���193), more maxillary teeth (11���13 vs. 9���10), and dorsal color pattern (dark uniform or stripes vs. blotches); from O. splendidus by having fewer scale rows at midbody (17 vs. 21), different supralabials entering the eye (SL 4���5 vs. 2���3), more infralabials (8 vs. 5), and shorter hemipenis (SC 13���14 vs. 19); from O. woodmasoni by having fewer ventrals (168���176 vs. 180���190) and more supralabials (8 vs. 6). By having deeply forked hemipenis without spines, O. condaoensis sp. nov. is similar to members of the O. cyclurus group (David et al. 2008a, Green et al. 2010, Neang et al. 2012, Smith 1943, Vassilieva 2015), which includes O. cattienensis Vassilieva, Geissler, Galoyan, Poyarkov, Van Devender & B��hme, O. chinensis (G��nther), O. cyclurus (Cantor), O. fasciolatus (G��nther), O. formosanus (G��nther). O. juglandifer (Wall), O. kampucheaensis Neang, Grismer & Dattry, O. kheriensis Acharji & Ray, O. macrurus (Angel), O. ocellatus (Morice), and O. saintgironsi David, Vogel & Pauwels. However, the new species can be distinguished from the following species by having 17 dorsal scale rows at midbody: O. cyclurus (19), O. fasciolatus (21), O. formosanus (19), O. juglandifer (19), O. kampucheaensis (15), O. khariensis (19), and O. ocellatus (19). It also differs from O. cyclurus by having smaller size (TL 552 mm vs. 710 mm), fewer dorsal scale rows on its neck (17 vs. 19), more maxillary teeth (11���13 vs. 9���10), arrangement of temporals (1+2 vs. 2+2), and dorsal color pattern (dark uniform or stripes vs. reticulate or narrow blotches); from O. fasciolatus by having smaller size (552 mm vs. 807 mm); fewer dorsal scale rows on the neck (17 vs. 21 or 23) and before the vent (15 vs. 17), more maxillary teeth (11���13 vs. 8���10), arrangement of temporals (1+2 vs. 2+2), and dorsal color pattern (dark uniform or stripes vs. reticulate or large blotches); from O. formosanus by having a smaller size (552 mm vs. 942 mm), fewer dorsal scale rows on the neck (17 vs. 19), fewer subcaudals (33���37 vs. 43���55) and infralabials (8 vs. 9), dorsal color pattern (dark uniform or stripes vs. reticulate) and hemipenis shape (no papillae vs. short papillae); from O. juglandifer by having fewer dorsal scale rows on the neck (17 vs. 19), fewer subcaudals (33���37 vs. 53���68), more supralabials (8 vs. 7), and dorsal color pattern (dark uniform or stripes vs. dark dots and cross-bars); from O. kampucheaensis by having more dorsal scale rows on the neck (17 vs. 15), fewer subcaudals (33���37 vs. 39), and dorsal color pattern (dark uniform or stripes vs. cross-bars); from O. khariensis by having fewer dorsal scale rows on the neck (17 vs. 19), fewer ventrals (168���176 vs. 196), and more maxillary teeth (11���13 vs. 8); and from O. ocellatus by having smaller size (552 mm vs. 852 mm), fewer dorsal scale rows on the neck (17 vs. 19), fewer infralabials (8 vs. 9 or 10), arrangement of temporals (1+2 vs. 2+2), and dorsal color pattern (dark uniform or stripes vs. reticulate or large blotches). Oligodon condaoensis sp. nov. differs from O. cattienensis by having more maxillary teeth (11���13 vs. 8���10), nasal (divided vs. entire), and dorsal color pattern (dark uniform or faint stripes vs. large blotches); from O. chinensis by having a smaller size (552 mm vs. 729 mm), more maxillary teeth (11���13 vs. 9���10), fewer subcaudals (33���37 vs. 47���64), and dorsal color pattern (dark uniform or faint stripes vs. black cross-bars); from O. macrurus by having more ventrals (168���176 vs. 143���152), fewer subcaudals (33���37 vs. 76���83), and much shorter hemipenis (SC 13���14 vs. 29); and from O. saintgoronsi by having a smaller size (552 mm vs. 676 mm), fewer dorsal scale rows on the neck (17 vs. 19), fewer subcaudals (33���37 vs. 53���59), much shorter hemipenis (SC 13���14 vs. 27���28), and dorsal color pattern (dark uniform or faint stripes vs. reticulate or large blotches). Table 3 compares the new species and other members of the O. cyclurus group. For members of the O. dorsalis group (Smith 1943, Vassilieva 2015), which includes O. catenatus (Blyth), O. dorsalis (Gray & Hardwicke), O. eberhardti Pellegrin, O. erythrogaster Boulenger, O. hamptoni Boulenger, O. lacroixi Angel & Bourret, and O. macdougalli Wall, O condaoensis sp. nov. differs from O. catenatus, O. dorsalis, O. eberhardti, O. erythrogaster, and O. hamptoni, by having deeply a forked hemipenis (vs. unforked or slightly forked) and an entire cloacal plate (vs. divided). It also differs from O. catenatus by having more dorsal scale rows at midbody (17 vs. 13), more maxillary teeth (11���13 vs. 7), and fewer ventrals (168���176 vs. 186���208); from O. dorsalis by having more dorsal scale rows at midbody (17 vs. 15) and more maxillary teeth (11���13 vs. 6���7); from O. eberhardti by having more dorsal scale rows (17���17���15 vs. 13���13���13), more supralabials (8 vs. 6), and fewer ventrals (168���176 vs. 189���190); from O. erythrogaster by having more maxillary teeth (11���13 vs. 7���8) and fewer subcaudals (33���37 vs. 42���59); and from O. hamptoni by having more scale rows at midbody (17 vs. 15), and more maxillary teeth (11���13 vs. 7) and more subralabials (8 vs. 5). The new species differs from O. lacroixi by having an entire cloacal plate (vs. divided), more scale rows at midbody (17 vs. 15), more supralabials (8 vs. 5), the presence of two internasals (vs. absent) and loreals (vs. absent); and from O. macdougalli by having an entire cloacal plate (vs. divided), the presence of loreals (vs. absent), more dorsal scale rows at midbody (17 vs. 13), fewer ventrals (168���176 vs. 200), and more supralabials (8 vs. 7). Oligodon condaoensis sp. nov. differs from members of the O. torquatus group (Green et al. 2010, Smith 1943, Vassilieva 2015), which includes O. cruentatus (G��nther), O. planiceps (Boulenger), O. theobaldi (G��nther), and O. torquatus (Boulenger), by having a deeply forked hemipenis (vs. unforked) without papillae (vs. presence) and an entire cloacal plate (vs. divided). It also differs from O. cruentatus by lacking spines on hemipenis (vs. presence) and having fewer maxillary teeth (11���13 vs. 14���16); from O. planiceps by lacking spines on hemipenis (vs. presence), more dorsal scale rows at midbody (17 vs. 13), more ventrals (168���176 vs. 132���145), more subcaudals (33���37 vs. 22���27), and more supralabials (8 vs. 4 or 5); from O. theobaldi by lacking spines on hemipenis (vs. presence) and having fewer maxillary teeth (11���13 vs. 15���16); and from O. torquatus by having more dorsal scale rows at midbody (17 vs. 15) and more ventrals (168���176 vs. 144���159), and fewer maxillary teeth (11���13 vs. 15���16). Oligodon condaoensis sp. nov. differs from other congeners in Indochina (Vietnam, Laos, Cambodia, Thailand, Myanmar, Malaysian Peninsular, and southern China [Yunnan]) as follows: from O. annamensis Leviton by having more dorsal scale rows (17���17���15 vs. 13���13���13), more supralabials (8 vs. 6), more infralabials (8 vs. 6), and more maxillary teeth (11���13 vs. 8), and by dorsal color pattern (dark uniform or faint stripes vs. cross-bars); from O. arenarius Vassilieva by having deeply forked hemipenis (vs. unforked), a larger size (552 mm vs. 389 mm), more maxillary teeth (11���13 vs. 6���8), and more ventrals (168���176 vs. 131���144); from O. booliati Leong & Grismer by having more supralabials (8 vs. 6 or 7), more ventrals (168���176 vs. 143���153), fewer subcaudals (33���37 vs. 54���60), and differen, Published as part of Nguyen, Sang Ngoc, Nguyen, Vu Dang Hoang, Le, Son Hong & Murphy, Robert W., 2016, A new species of kukri snake (Squamata: Colubridae: Oligodon Fitzinger, 1826) from Con Dao Islands, southern Vietnam, pp. 261-273 in Zootaxa 4139 (2) on pages 263-270, DOI: 10.11646/zootaxa.4139.2.9, http://zenodo.org/record/256798, {"references":["David, P., Vogel, G. & Pauwels, O. S. G. (2008 a) A new species of the genus Oligodon Fitzinger, 1826 (Squamata: Colubridae) from southern Vietnam and Cambodia. Zootaxa, 1939, 19 - 37.","David, P., Vogel, G. & Rooijen, J. V. (2008 b) A revision of the Oligodon taeniatus (Gunther, 1861) group (Squamata: Colubridae), with the description of three new species from the Indochinese region. Zootaxa, 1965, 1 - 49.","Green, M. D., Orlov, N. L. & Murphy, R. W. (2010) Toward a phylogeny of the kukri snakes, genus Oligodon. Asian Herpetological Research, 1, 1 - 21.","Neang, T., Grismer, L. L. & Daltry, J. C. (2012) A new species of kukri snake (Colubridae: Oligodon Fitzinger, 1826) from the Phnom Samkos Wildlife Sanctuary, Cardamom Mountains, southwest Cambodia. Zootaxa, 3388, 41 - 55.","Smith, M. A. (1943) The Fauna of British India, Ceylon and Burma, including the whole of the Indo-Chinese subregion. Reptiles and Amphibians. Vol. III. Serpentes. Taylor and Francis Ltd, London, 525 pp.","Vassilieva, A. B. (2015) A new species of the genus Oligodon Fitzinger, 1826 (Squamata: Colubridae) from coastal southern Vietnam. Zootaxa, 4058 (2), 211 - 226. http: // dx. doi. org / 10.11646 / zootaxa. 4058.2.4","David, P., Nguyen, T. Q., Nguyen, T. T., Jiang, K., Chen, T., Teynie, A. & Ziegler, T. (2012) A new species of the genus Oligodon Fitzinger, 1826 (Squamata: Colubridae) from northern Vietnam, southern China and central Laos. Zootaxa, 3498, 45 - 62.","Campden-Main, S. M. (1970) A field guide to the snakes of South Vietnam. Smithsonian Institution, Washington, 114 pp."]}

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

Bryson, Robert W., Linkem, Charles W., Dorcas, Michael E., Lathrop, Amy, Jones, Jason M., Alvarado-Díaz, Javier, Grünwald, Christoph I., Murphy, Robert W. (2014): Multilocus species delimitation in the Crotalus triseriatus species group (Serpentes: Viperidae: Crotalinae), with the description of two new species. Zootaxa 3826 (3): 475-496, DOI: http://dx.doi.org/10.11646/zootaxa.3826.3.3

Crotalus campbelli sp. nov. Figs. 7���8, Table 3 Crotalus triseriatus ��� Boulenger (1896): 581 (in part). Crotalus triseriatus armstrongi ��� Dorcas (1992): 87 (in part). Crotalus triseriatus armstrongi ��� Bryson et al. (2011): 699 (in part). Crotalus triseriatus armstrongi ��� Reyes-Velasco et al. (2009): 118. Crotalus armstrongi ��� Reyes-Velasco et al. (2013): 528 (in part). Holotype. Adult female (KU 73649) collected 25 October 1962 by Percy L. Clifton (field number PLC 3216) in the Sierra de Cuale, 9 km N El Teosinte, municipality of Talpa de Allende, state of Jalisco, Mexico. Paratypes. 5 specimens. Mexico: JALISCO: same collection data as holotype (KU 73650). Las Playitas, Las Joyas, Sierra de Manantl��n, municipality of Autl��n de Navarro; collected September 1985 by E. Fanti-Echegoyen (UTA R- 16352). Las Joyas, Sierra de Manantl��n, municipality of Autl��n de Navarro; collected September 1985 by E. Fanti-Echegoyen (UTA R- 16353). ca. 25 km SE Autl��n, ca. 2.1 km (by dirt road) SE Manantl��n; collected 27 July 1975 by G. M. Tilger and R. G. Arndt (AMNH R- 113191). Lago de Juanacatl��n, Sierra de Mascota, municipality of Mascota (20 �� 37 ' 30.94 "N, 104 �� 43 ' 36.30 "W; 2009 m asl; WGS 84); collected 10 April 2011 by R. W. Bryson Jr. and M. Torocco (MZFC 28669). Diagnosis. Crotalus campbelli can be distinguished from all members of the C. triseriatus species group except C. armstrongi by the combination of the following characters: (1) presence of intercanthals, (2) infrequently divided upper preocular (9.1 % of the time), (3) 150���154 ventrals in males, 147���152 in females, (4) 31���32 subcaudals in males, 22���26 in females, (5) small rattle (proximal rattle width 11.0��� 14.6 % of head length), (6) long tail (9.1 ���11.0% of total body length in males, 7.5���8.9 % in females), (7) pale interspaces between dorsal and lateral blotches, (8) heavy venter mottling, (9) dark proximal rattle and underside of tail, and (10) usually a single large anterior intercanthal. Crotalus campbelli can be distinguished from C. armstrongi based on higher mean number of ventrals (152 in males and 149 in females vs. 141 and 144), higher mean number of subcaudals in males (31 vs. 28), less frequently divided upper preocular (9.1 % vs. 14.3 %), proportionately longer tail in males (10.3 % of total body length vs. 9.7 %), smaller mean proximal rattle width (13.0% of head length vs. 14.0%), higher mean number of dorsal blotches (48 vs. 42), and higher number of tail bands (mode of 9 vs. 6). Crotalus campbelli is most similar to members of the C. triseriatus group distributed along the Trans-Mexican Volcanic Belt, including C. pusillus, C. armstrongi, C. triseriatus, and C. tlaloci. Crotalus campbelli is distinguished from C. pusillus by possessing intercanthals and an infrequently divided upper preocular, from C. tlaloci by having an infrequently divided upper preocular, variable number of intercanthals, fewer ventrals (in females, 147���152 vs. 156���165; in males, mean number 152 vs. 156), lower mean number of subcaudals in females (24 vs. 28), proportionately shorter tail in females (8.2 % of total length vs. 9.2 %), and higher mean number of dorsal blotches (48 vs. 43), and from C. triseriatus by a higher number of ventrals (in males, 150���154 vs. 134���146; in females, mean number higher: 149 vs. 142), higher mean number of subcaudals in males (31 vs. 28), proportionately smaller proximal rattle (13.0% of head length vs. 15.8 %), and by having pale interspaces between dorsal and lateral blotches. Crotalus campbelli is most similar in general appearance to C. armstrongi, but can be distinguished from this species by characters mentioned above. Half of the specimens of C. campbelli also possess a single, large anterior intercanthal. This scale arrangement is rarely seen in C. armstrongi and C. triseriatus. Crotalus campbelli is easily distinguished from C. ravus by the lack of large head plates in the parietal region. Description of the holotype. Rostral broader than high (3.7 x 2.6 mm); two internasals, in medial contact, slightly wider than long, convex through center of scale; two canthals, large, convex, circular, separated by a single, large, convex intercanthal bordered anteriorly by three small scales; four large intersupraoculars posterior to intercanthals, followed by multiple rows of small intersupraoculars. Nostril centered between prenasal and postnasal scales, prenasal larger than postnasal and wrapping around anterior aspect of rostrum; single loreal. Loreal pit midway between eye and naris, below line from middle of eye to naris, bordered by single prelacunal, postlacunal, lower preocular; prelacunal contacting second and third supralabials and two prefovials; three prefovials; single postfovial contacting postlacunal, lower preocular, first subocular, and fourth supralabial. Two preoculars, upper large and convex, contacting supraocular and canthal, lower preocular thin and long; three suboculars, anterior largest and in contact with fourth supralabial; three interoculabials posterior to anterior subocular; two postoculars, upper twice as large as lower. Supralabials 13 / 13; infralabials 11 / 12; first infralabials in medial contact posterior to triangular mental; genials together resemble wings. Midbody dorsal scale rows 23���25; preventral single; ventrals 152; subcaudals 22, nondivided; eight rattle fringe scales; tail bearing three rattle segments. Ground color, in preservative, light brown with 54 dark dorsal body blotches, irregularly edged in black with a very thin light outer edge on most blotches, blotches wider than long, interspaces between blotches 1���2 scales long; smaller, vertically elongated auxiliary blotches evident laterally below the dorsal blotches, often separated from the dorsal blotches by white scales. Head marked with parallel rows of small irregular blotches across prefrontals, supraoculars, and much of the occipital area, terminated by paired occipital blotches; supralabials and infralabials ground color white, stippled with dark brown spots; distinct dark postocular stripe, uniform in width, extending from posterior of eye to above the angle of the mouth and then downwards to jawline; postocular stripe bordered by a darker band on both dorsal and ventral margins, dorsally a narrow region of light brown extends from the posterior axis of the eye along the dorsal margin of the postocular band, ventral to the postocular band is a region of white with brown stippling, supralabial 8 is nearly completely white, with moderate stippling but no postocular band, supralabials 8���13 are divided by the postocular band; two large dark brown blotches are on the borders of supralabials 5���7; scale margins of infralabials with dark brown regions making for a series of dark bands, bands do not extend beyond infralabials; gular scales stippled with no pattern; ventral scutes stippled, more heavily on posterior half of scales, with total stippling intensifying past midbody; distal 1 / 3 ventral scutes become dark brown with a lighter band along the midline, continuing to the vent. Subcaudal scales dark brown to black; eight dark brown tail bands; proximal rattle black, distal sections brown. Color in life. Color in life within the type series is only known for one paratype (MZFC 28669), shown in Fig. 8. Variation. Three of the six specimens lack a single, large anterior intercanthal. The holotype, KU 73649, has a single large intercanthal bordered anteriorly by three small, seemingly anomalous scales (Fig. 7). Similarly, UTA R- 16353 has paired anterior intercanthals separated by a small but elongated scale. The third specimen, AMNH R- 113191, has paired anterior intercanthals. The single juvenile (AMNH R- 113191) has a cream-colored proximal rattle and pale-colored tail ventrally. Variation in meristic, morphometric, and color pattern characters within the type series is listed in Table 3. Etymology. The specific epithet is a patronym honoring Jonathan A. Campbell for his many years of field research on Mexican rattlesnakes and for his decades of unwavering support to students of Mexican herpetology. Habitat and distribution. Crotalus campbelli is found in rocky open breaks within montane forest (Fig. 9) along the far western regions of the Trans-Mexican Volcanic Belt. Much of this forest is covered with remnant patches of cloud forest (Ponce-Reyes et al. 2012). This species is known from western Jalisco and the Sierra de Manantl��n in southern Jalisco/northwestern Colima. A narrow low-elevation valley appears to separate the range of C. campbelli from C. armstrongi to the east (Fig. 3)., Published as part of Bryson, Robert W., Linkem, Charles W., Dorcas, Michael E., Lathrop, Amy, Jones, Jason M., Alvarado-D��az, Javier, Gr��nwald, Christoph I. & Murphy, Robert W., 2014, Multilocus species delimitation in the Crotalus triseriatus species group (Serpentes: Viperidae: Crotalinae), with the description of two new species, pp. 475-496 in Zootaxa 3826 (3) on pages 486-489, DOI: 10.11646/zootaxa.3826.3.3, http://zenodo.org/record/225469, {"references":["Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History). Vol. III. Trustees of the British Museum, London, xiv + 727 pp., 15 plates.","Dorcas, M. E. (1992) Relationships among montane populations of Crotalus lepidus and Crotalus triseriatus. In: Campbell, J. A. & Brodie Jr., E. D. (Eds.), Biology of the Pitvipers. Selva, Tyler, Texas, pp. 71 - 88.","Bryson, R. W., Murphy, R. W., Lathrop, A. & Lazcano-Villareal, D. (2011) Evolutionary drivers of phylogeographical diversity in the highlands of Mexico: a case study of the Crotalus triseriatus species group of montane rattlesnakes. Journal of Biogeography, 38, 697 - 710. http: // dx. doi. org / 10.1111 / j. 1365 - 2699.2010.02431. x","Reyes-Velasco, J., Hermosillo-Lopez, I. A., Grunwald, C. I. & Avila-Lopez, O. A. (2009) New state records for amphibians and reptiles from Colima, Mexico. Herpetological Review, 40, 117 - 120.","Reyes-Velasco, J., Meik, J. M., Smith, E. N. & Castoe, T. A. (2013) Phylogenetic relationships of the enigmatic longtailed rattlesnakes (Crotalus ericsmithi, C. lannomi, and C. stejnegeri). Molecular Phylogenetics and Evolution, 69, 524 - 534. http: // dx. doi. org / 10.1016 / j. ympev. 2013.07.025","Ponce-Reyes, R., Reynoso-Rosales, V. - H., Watson, J. E. M., VanDerWal, J., Fuller, R. A., Pressey, R. L. & Possingham, H. P. (2012) Vulnerability of cloud forest reserves in Mexico to climate change. Nature Climate Change, 2, 448 - 452."]}

Nguyen, Sang Ngoc, Yang, Jun-Xiao, Le, Thanh-Ngan Thi, Nguyen, Luan Thanh, Orlov, Nikolai L., Hoang, Chung Van, Nguyen, Truong Quang, Jin, Jie-Qiong, Rao, Ding-Qi, Hoang, Thao Ngoc, Che, Jing, Murphy, Robert W., Zhang, Ya-Ping (2014): DNA barcoding of Vietnamese bent-toed geckos (Squamata: Gekkonidae: Cyrtodactylus) and the description of a new species. Zootaxa 3784 (1), DOI: http://dx.doi.org/10.11646/zootaxa.3784.1.2

Crotalus tlaloci sp. nov. Figs. 4–5, Table 3 Crotalus triseriatus triseriatus — Campbell & Lamar (2004): 527 (Map 90), 593 (in part). Crotalus triseriatus triseriatus — Flores-Villela & Hernández-García (1989): 16. Crotalus triseriatus triseriatus — Pérez-Ramos et al. (2000): 34. Crotalus triseriatus triseriatus — Flores-Villela & Hernández-García (2006): 270. Holotype. Adult female (MZFC 3666) collected 20 June 1986 by Efrain Hernández-García in “Los Llanos” (18 ° 36 ’N, 99 ° 37 ’W; 2200–2300 m above sea level; asl hereafter), 10 km by road from Taxco to Tetipac, Sierra de Taxco, municipality of Tetipac, state of Guerrero, Mexico. Paratypes. 11 specimens. MEXICO: GUERRERO: Cerro del Huizteco, Sierra de Taxco, municipality of Taxco (18 ° 36 'N, 99 ° 36 'W; 2300–2520 m asl); collected 22–23 August 1986 by E. Hernández-García (MZFC 3664–3665). “Arroyo las Damas”, Sierra de Taxco, municipality of Tetipac (18 ° 38 'N, 99 ° 37 'W; 1600–1850 m asl); collected by E. Hernández-García (MZFC 3666). ESTADO DE MÉXICO: Acatitlán, municipality of Valle de Bravo; collected 7 September 1988 by T. Hentschel-Maida (MZFC 4324). Los Álamos, municipality of Valle de Bravo (19 ° 11 ' 20.2 "N, 100 °03' 57.2 "; 2201 m asl; NAD 27 Mexico); collected 23 May 2008 by J. Jones, C. I. Grünwald, and R. W. Bryson Jr. (HINIRENA 725–726). Los Álamos, municipality of Valle de Bravo (19 ° 11 ' 20.2 "N, 100 °03' 57.2 "; 2201 m asl; NAD 27 Mexico); collected 22 July 2009 by R. W. Bryson Jr. and M. Torocco (MZFC 25114–25115). MORELOS: Km 12, Carr. Cuernavaca-Ocuilán, municipality of Cuernavaca; collected 17 March 1990 by M. Torres Chávez (MZFC 4657). Carr. Cuernavaca-Ocuilán, near state border, municipality of Cuernavaca (18 ° 58 ' 54.43 "N, 99 ° 18 ' 20.43 "W; 2268 m asl; WGS 84); collected 13 June 2009 by J. Jones, C. I. Grünwald, and R. W. Bryson Jr. (MZFC 25111). MICHOACÁN: N Arroyo Seco, municipality of Aporo (19 ° 40 ' 28.3 "N, 100 ° 22 ' 35.8 "W; 2463 m asl; NAD 27 Mexico); collected 24 May 2008 by J. Jones, C. I. Grünwald, and R. W. Bryson Jr. (HINIRENA 724). Diagnosis. Crotalus tlaloci can be distinguished from all members of the C. triseriatus species group by the combination of the following characters: (1) presence of intercanthals, (2) undivided upper preocular, (3) 152–164 ventrals in males, 156–165 in females, (4) 27–33 subcaudals in males, 22–32 in females, (5) small rattle (proximal rattle width 11.1–14.5 % of head length), (6) long tail (8.9–11.3 % of total body length in males, 8.0– 10.7 % in females), (7) usually two pairs of symmetrical, similarly sized intercanthals, and (8) dark postocular stripe that noticeably narrows before reaching the posterior of the eye. Crotalus tlaloci is most similar to species of the C. triseriatus group distributed along the Trans-Mexican Volcanic Belt. Crotalus tlaloci is distinguished from these species by the presence of intercanthals (lacking in C. pusillus), undivided upper preocular (divided 14.3 % of the time in C. armstrongi and 9.4 % of the time in C. triseriatus), high number of ventrals (overlapping with C. pusillus, but mean number in C. tlaloci higher than C. pusillus: 156 vs. 154, respectively), high number of subcaudals (mean number in females higher than in C. pusillus, C. armstrongi, and C. triseriatus: 28 vs. 26, 24, and 24; in males, higher than in C. armstrongi and C. triseriatus: 30 vs. 28 in both), proportionately small proximal rattle (mean width smaller than in C. armstrongi and C. triseriatus: 12.8 % of head length vs. 14.0% and 15.8 %), and proportionately longer tail (mean length in females higher than in C. pusillus, C. armstrongi, and C. triseriatus: 9.2 % of total length vs. 8.6 %, 8.3 %, and 8.3 %). Most specimens (10 / 12) possess two pairs of symmetrical, similarly sized intercanthals, creating the appearance of butterfly wings. Of the 100 specimens in the C. triseriatus group that we examined, this symmetrical paired arrangement of intercanthal scales in the prefrontal region (“butterfly wings”) was observed in only one other specimen (C. armstrongi, CNAR 4498). Crotalus tlaloci also possess a dark postocular stripe that noticeably narrows before reaching the posterior of the eye. In C. pusillus, C. armstrongi, and C. triseriatus, the postocular stripe is generally of uniform width, although on rare occasions in C. pusillus and C. armstrongi it tapers slightly before reaching the eye. Description of the holotype. Rostral broader than high (4.0 x 2.8 mm); two internasals, in medial contact, wider than long, rectangular, convex through center of scale; two canthals, large, convex, separated by two pairs of square intercanthals; four large intersupraoculars posterior to intercanthals, followed by multiple rows of small intersupraoculars. Naris centered between prenasal and postnasal scales, prenasal larger than postnasal and wrapping around anterior aspect of rostrum; two loreals, lower larger; small upper loreal between canthal, internasal, postnasal, and lower loreal. Loreal pit midway between eye and naris, below line from middle of eye to naris, bordered by single prelacunal, postlacunal, lower preocular, and lower loreal; prelacunal contacting second and third supralabials and a single prefovial (two prefovials on left); two prefovials on right, three on left; single postfovial contacting postlacunal, lower preocular, first subocular, and fourth supralabial. Two preoculars, upper large and convex, contacting supraocular and canthal, lower preocular thin and long; three suboculars, anterior largest and in contact with fourth supralabial; three interoculabials posterior to anterior subocular; two postoculars, dorsal twice as large as ventral. Supralabials 12 / 12; infralabials 13 / 13; first infralabials in medial contact posterior to triangular mental; genials together resemble a heart shape. Midbody dorsal scale rows 22–23; preventral single; ventrals 162; subcaudals 27, last subcaudal row divided into three scales; nine rattle fringe scales; tail bearing four rattle segments. Coloration significantly faded in preservative with body blotches and bands difficult to discern from ground color. Ground color gray; occasional black speckling along body. Blotches faded to ground coloration in many places, approximately 38 blotches visible. Blotches bordered by lighter coloration than ground color, also heavily faded. Ground color of head gray, heavily stippled with dark brown throughout dorsal and anterior lateral regions of the head; rostral and supralabials 1–4 heavily stippled, supralabials 5–7 with lighter stippling in the scale center; infralabials with brown stippling on scale margins with white scale centers, extending to mouth; gular scales lightly stippled, decreasing in frequency towards the midline of the head. Ventral scutes evenly stippled, increasing in intensity posterior to mid-body with scutes becoming almost completely brown by the tail. Six dark tail bands, the first two bordered by white bands; caudal scales dark brown. Proximal rattle black, distal sections brownish. Color in life. Color in life varies (Fig. 5), with most specimens vibrantly colored, although one is darkly pigmented (MZFC 25111; Fig. 5). The holotype is known only from the preserved specimen. Variation. Two specimens lack symmetrical paired intercanthals. The first, HINIRENA 725, has one pair of anterior intercanthals but only a single large posterior intercanthal. The second, HINIRENA 724, has one pair of anterior intercanthals, one of which is subdivided, followed by a single large posterior intercanthal. Supralabials in five specimens (MZFC 3664, 3665, 4657, 25111, 25114) are horizontally divided. This split was in the 7 th, 8 th, or 9 th supralabial and when present, occurred in supralabials on both sides of the head. Juveniles (MZFC 3664, 3665, 3667, HINIRENA 726) have cream-colored proximal rattles and pale-colored tails ventrally, typical of juveniles in other species in the C. triseriatus group. Variation in meristic, morphometric, and color pattern characters within the type series is listed in Table 3. Etymology. This species is named for Tláloc, the Aztec god of rain. Habitat and distribution. Crotalus tlaloci inhabits open areas in cloud forest and humid oak-pine forest along the lower slopes of the Trans-Mexican Volcanic Belt. Although one record in the Sierra de Taxco (“Arroyo las Damas”) is at 1850 m (Flores-Villela & Sánchez-H 2003), most records are at around 2000–2400 m asl. This species is known from the states of Guerrero, Estado de México, Michoacán, and Morelos, and may range into western Puebla. The vegetation where C. tlaloci is found is characterized by broad-leaf oaks, such as Quercus candicans and Q. laurina, and dense undergrowth (Fig. 6), and is distinctly different than the drier pine-oak forest inhabited by C. triseriatus. The distribution of C. tlaloci overlaps the ranges of two alligator lizards, Barisia herrerae Zaldívar-Riverón & Nieto-Montes de Oca 2002 and B. rudicollis (Wiegmann 1828) (Zaldívar-Riverón & Nieto-Montes de Oca 2002). Interestingly, both of these alligator lizards occur in similar humid forest habitat at elevations of 2000–2500 m asl, and appear ecologically isolated from B. imbricata (Wiegmann 1828), which inhabits the surrounding drier pine-oak forest (Zaldívar-Riverón & Nieto-Montes de Oca 2001, 2002). Specimens of C. tlaloci are generally found in rocky open forest breaks and edges of cloud or humid oak-pine forest. However, we found an adult gravid female and juvenile (HINIRENA 725, 726) under logs in a clearing relatively devoid of rocky habitat.

Cyrtodactylus puhuensis sp. nov. P�� Hu bent-toed gecko (Figs. 3���6) Holotype. KIZ 11665, an adult male from P�� Hu Nature Reserve, Thanh Hoa Province, Vietnam, 20 �� 33 ��� N, 104 �� 53 ��� E, 638 m, collected on 21 August 2011 by Hoang Van Chung, Yang Jun Xiao and Nguyen Ngoc Sang. Diagnosis. Medium sized, snout-vent length 79.24 mm; body moderately robust; tail 82.59 mm, longer than snout-vent length, with a series of medium enlarged transversal subcaudals; a series of medium enlarged femoral scales continue from precloacal scales; femoral scales without pores; males with five precloacal pores; supralabials 8; infralabials 10; narrow subdigital lamellae on fourth toe 23; ventral scales 36; and dorsal tubercles smooth or slightly keeled. Description of holotype. In addition to diagnosis. Head relatively depressed, height 9.53 mm, width 15.00 mm (height/width ratio 0.64), distinct from neck; supralabials 8, continuous with a row of small scales extending to the corner of the mouth, first contacting nostril; infralabials 10; supra- and infralabials nearest to rostral and mental largest in size; rostral large, width 3.27 mm, height 2.40 mm, vertical sunken groove in middle, contacting nostril; two internasals contact each other; mental nearly triangular, posteriorly contacting two large postmentals that contact one another and first infralabials (Fig. 5 C���E); gular scales much smaller than ventrals; head scales small, granular, with scattered tubercles in occipital and temporal areas; eye relatively large, 5.85 mm (eye/head width ratio 0.61), pupil vertical; supraciliary scales large, cover the eye anteriorly, superiorly, and posteriorly; ear opening oval, 1.90 mm, obliquely oriented, much smaller than eyes (ear/eye ratio 0.32); eye-nostril distance 7.00 mm, eyesnout distance 8.83 mm, eye-ear distance 6.27 mm, distance between eyes 8.06 mm, distance between nostrils 2.40 mm. Body elongate, trunk length 35.73 mm, with weak ventral-lateral folds; dorsal scales small, granular, not homogeneous; dorsal and lateral tubercles round, smooth or slightly keeled, not in rows; dorsal tubercles between thighs and base of tail conical and keeled; ventral scales fairly large, smooth, and flattened, about 36 midbody transversal ventrals; dorsal surface of limbs with granular scales and slightly keeled conical tubercles; basal subdigital lamellae narrow and irregular, largest in joints of digits; subdigital lamellae well developed at distal phalanx, as broad as the digit. Subdigital lamellae under fourth finger 18 and toe 23; digits unwebbed; forearm length 13.40 mm, crus length 16.00 mm. Precloacal scales large, flattened, and imbricate; about 8 large imbricate scales under a series of 8 precloacal scales arranged in ��-shape that bears five precloacal scales with pores; a series of moderately enlarged femoral scales on each side of thigh extend from the eight precloacal scales (Fig. 5 A); precloacal groove and femoral pores absent; tail round in cross-section, 82.59 mm in length, longer than snout-vent length; transversal subcaudals medium size, about a half size of tail width (Fig. 5 B); dorsum of proximal tail with conical tubercles extending down to the base tail. Color in life (Fig. 4). Dorsal and lateral background of body chocolate-black; dorsum with faint and irregular pale-yellowish transverse bands; dorsal tubercles at the transverse bands yellow to white, others colored as background; flanks with yellow to white tubercles; head dark brown with yellowish pattern; eyelids yellow; eye brown, with black vertical pupil; nuchal band chocolate-black, interrupted at nape; upper surface of legs and digits dark brown with yellow to white bands or blotches; unregenerated tail with seven black and white to yellowish alternating rings; white rings much more narrow than black; lower surfaces of flanks and legs white. Color in preservative (Fig. 6 A, B). In preservative, color slightly faded and pattern unchanged; body background black or dark brown, dorsal transverse bands white; dorsum and flanks with white tubercles. DNA barcode: GenBank Accession No. KF 929529. Comparisons. The new species was compared to other congeners in Indochina by using the following morphological characters: precloacal pores, subcaudal scales, ventral scales, continuous medium enlarged femoral scales, dorsal tubercles, dorsal pattern, nuchal band, and number of digital lamellae under the 4 th toe. Cyrtodactylus puhuensis sp. nov. differs from Vietnamese congeners, from C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler 2012, C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler 2012, C. cattienensis Geissler, Nazarov, Orlov, B��hme, Phung, Nguyen & Ziegler 2009, C. cryptus Heidrich, R��sler, Vu, B��hme & Ziegler 2007, C. dati Ngo 2013, C. huynhi Ngo & Bauer 2008, C. irregularis (Smith 1921 b, 1935), C. pseudoquadrivirgatus R��sler, Nguyen, Vu, Ngo & Ziegler 2008, and C. ziegleri Nazarov, Orlov, Nguyen & Ho 2008, in having a series of moderately enlarged subcaudal scales and continuous precloacal-femoral scales. The new species differs from C. martini Ngo 2011 in having a series of transversal and enlarged subcaudal scales (versus small or medium ones, not transversely enlarged) and a higher number of precloacal pores (5 versus 4 in all four males from the type series). By having series of moderately enlarged femoral scales continuous to the precloacal scales, C. puhuensis sp. nov. differs from C. badenensis Nguyen, Orlov & Darevsky 2006, C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen 2007, C. cucphuongensis Ngo & Onn 2011, C. eisenmanae Ngo 2008, C. grismeri Ngo 2008, C. huongsonensis Luu, Nguyen, Do & Ziegler 2011, C. kingsadai Ziegler, Phung, Le & Nguyen 2013, C. nigriocularis Nguyen, Orlov & Darevsky 2006, and C. takouensis Ngo & Bauer 2008. In addition, the new species differs from C. badenensis, C. cucphuongensis, C. eisenmanae, C. grismeri, and C. nigriocularis in having 5 precloacal pores (versus 0, or 0���2 in the last species); from C. huongsonensis and C. kingsadai by absence of femoral pores (versus 17 or 3���7 femoral pores on both thighs, respectively); and from C. caovansungi and C. bichnganae Ngo & Grismer 2010 by lacking femoral pores (versus 3, or 9 on each thigh, respectively) and having a less precloacal pores (5 versus 9 or 10, respectively). Our examination of the type, paratype and additional specimens collected from the type locality (Appendix 1) showed C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau 2007 differed from the new species. First, C. chauquangensis has a uniform transversal butterfly-shaped pattern on dorsum (Fig. 7) while the new species shows faint, thin, and irregular dorsal bands. Second, femoral scale rows in C. chauquangensis are undeveloped; they are the same size as other femoral scales. The new species has series of moderately enlarged femoral scales (Fig. 5 A). Third, C. chauquangensis has 6���7 precloacal pores, except for topotype CQ 526, which has 8 pores. In contrast, the new species has 5 pores only. The new gecko differs from C. condorensis (Smith 1921 a) by the distribution of precloacal pores (in a chevron versus 4���7 in two juxtaposed rows) and dorsal tubercles (smooth or slightly keeled versus subtrihedral) (Smith 1935; Ngo & Grismer 2012); from C. paradoxus (Darevsky & Szczerbak 1997) by having more and differently arranged precloacal pores (5 in chevron versus 0���4 in two juxtaposed rows) (Orlov et al. 2007); and from C. thochuensis Ngo & Grismer 2012 by the chevron arrangement of its precloacal pores (versus in two juxtaposed rows), smooth or slightly keeled dorsal tubercles (versus strongly keeled), normal sized scales on heel (versus enlarged), and having the dorsal caudal tubercles extending to the base of the tail only (versus to about 1 / 3 the length of the tail). Cyrtodactylus puhuensis sp. nov. differs from C. hontreensis Ngo, Grismer & Grismer 2008, C. intermedius (Smith 1917, 1935), and C. phuquocensis Ngo, Grismer & Grismer 2010 in having faint, irregular pale yellowish dorsal bands. In contrast, C. hontreensis has 3 broad bands between limbs and C. intermedius and C. phuquocensis have 4. The new species also differs in its absence of nuchal band, which is always present in the other species, and in having 5 precloacal pores versus 7���8 in C. hontreensis, 8���10 in C. intermedius and 6���8 in C. phuquocensis. The new species differs from C. phongnhakebangensis Ziegler, R��sler, Herrmann & Vu 2003 and C. roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz 2010 in lacking femoral pores (versus 27���42 and 20��� 28, respectively, precloaacal-femoral pores situated as one continuous row). The new gecko differs from C. yangbayensis Ngo & Onn 2010 in possessing fewer precloacal pores (5 versus 6���8), more subdigital lamellae under the fourth toe (23 versus 15���17), medium sized scales on heel (versus 9���10 enlarged scales), and seven pale rings on the tail (versus 9���11 irregular bands). For Laotian congeners, Cyrtodactylus puhuensis sp. nov. differs from C. buchardi David, Teyni�� & Ohler 2004, C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler 2011, and C. wayakonei Nguyen, Kingsada, R��sler, Auer & Ziegler 2010 in having a series of enlarged femoral scales (versus without enlarged femoral scales). The new species also differs from C. buchardi and C. wayakonei in having enlarged transversal subcaudal scales (versus small or medium scales, not transversally enlarged); and from C. pageli in lacking a nuchal band (versus dark nuchal band). Cyrtodactylus puhuensis sp. nov. differs from C. jarujini Ulber 1993 and C. lomyenensis Ngo & Pauwels 2010 in lacking femoral pores (52���54 and 39���40, respectively, continuous precloacal-femoral pores). The new gecko differs from C. interdigitalis Ulber 1993 and C. teyniei David, Nguyen, Schneider & Ziegler 2011 in having fewer precloacal pores (5 versus 14) and also differs from the former by lacking femoral pores (versus 8���9). From Myanmar congeners, Cyrtodactylus puhuensis sp. nov. differs from C. ayeyarwadyensis Bauer 2003, C. brevidactylus Bauer 2003, C. gansi Bauer 2003, C. mandalayensis Mahony 2009, C. tamaiensis Mahony 2009, and C. wakeorum Bauer 2003, in having moderately enlarged transversal subcaudal scales. The new species also differs from C. ayeyarwadyensis, C, brevidactylus, and C. gansi in having few precloacal pores (versus 10 ���28, 8, and 16��� 29, respectively); from C. gansi in lacking a shallow precloacal groove in males; and from C. tamaiensis in lacking femoral pores (versus 40 precloacal-femoral pores in an almost continuous series, Mahony 2009). From C. pulchellus Gray and C. rubidus (Blyth 1861) the new species differs in lacking a precloacal groove (Smith 1935; Taylor 1963). Cyrtodactylus puhuensis sp. nov. differs from C. aequalis Bauer 2003, in lacking femoral pores (versus 3���4 minute femoral pores) and possesing more ventral scales (36 versus 24); from C. annandalei Bauer 2003, in having less precloacal pores (5 versus 11���12), lacking femoral pores (versus 10���11), and having a dorsal pattern of faint and irregular pale yellowish bands (versus six dark bands); from C. chrysopylos Bauer 2003, in having less precloacal pores (5 versus 10 and a single, pored scale posterior to precloacal series); from C. consobrinoides (Annandale), in having more ventral scales (36 versus 24���30), lacking a nuchal band, and possessing a dorsal pattern of faint and irregular pale yellowish bands (versus 6���7 narrow dark cross-bars or transverse markings, Smith 1935); from C. feae (Boulenger 1893), in lacking femoral pores (Bauer 2003), and having a dorsal pattern of faint and irregular pale yellowish bands (versus four dark cross-bars, Smith 1935); from C. oldhami (Theoblad 1876), in having more precloacal pores (5 versus 0���4), absence of a nuchal band, and a having a dorsal pattern of faint and irregular pale yellowish bands versus white spots) (Smith 1935); from C. peguensis (Boulenger 1893), in having smaller number of precloacal pores (5 versus 7���9) and continuous series of precloacal-femoral scales (Smith 1935; Taylor 1963); from C. russelli Bauer 2003 and C. slowinskii Bauer 2003, in having less precloacal pores (5 versus 15 and 9, respectively) and lacking femoral pores (versus 16���19 and 11 pores on each thigh, respectively); and from C. variegatus (Blyth 1859), in having more ventral scales (36 versus 22) and lacking femoral pores (versus 32 continuous precloacal-femoral pores (Smith 1935). For Thai congeners, Cyrtodactylus puhuensis sp. nov. differs from C. quadrivirgatus Taylor 1962 and C. angularis (Smith 1921 b), in having moderately enlarged subcaudal scales (versus small; Smith 1935; Taylor 1963). The new gecko differs from C. phuketensis Sumontha, Pauwels, Kunya, Nitikul, Samphanthamit & Grismer 2012, in lacking a precloacal groove and femoral pores (versus 33���36 continuous precloacal-femoral pores). The new species differs from the following species in lacking femoral pores: C. astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels 2012, C. auribalteatus Sumontha, Panitvong & Deein 2010, C. brevipalmatus (Smith 1923), C. chanhomeae Bauer, Sumontha & Pauwels 2003, C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya 2010, C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya 2009, C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels 2012, C. marmoratus (Gray 1831), and C. trigroides Bauer, Sumontha & Pauwels 2003 (Smith 1935; Taylor 1963). Cyrtodactylus puhuensis sp. nov. can be distinguished from C. papilionoides Ulber & Grossmann 1991, in having a dorsal pattern of faint and irregular pale yellowish bands (versus serrated, broad, dark bands with pale margins or transversal rows of blotches), and having a continuous series of 36 digital lamellae under the 4 th toes (versus an interrupted series of 12���16 lamellae; Heidrick et al. 2007; Nguyen et al. 2010); from C. consobrinus (Peters 1871), in having less precloacal pores (5 versus 9���11) and a dorsal pattern of faint and irregular pale yellowish bands (versus 8 dark, transverse cross-bands, Boulenger 1885); from C. sanook Pauwels, Sumontha, Latinne & Grismer 2013, in having medium enlarged femoral scale rows (versus well-developed); from C. sumonthai Bauer, Pauwels & Chanhome 2002, in having more precloacal pores (5 versus 2 minute pores), and a series of moderately enlarged femoral scales (versus the absence of enlarged femoral scales); from C. surin Chan-ard & Makchai 2011, in having more ventral scales (36 versus 25) and smooth or slightly keeled dorsal tubercles (versus very large, trihedral, and keeled); and from C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome 2004, in having fewer precloacal pores (5 versus 8���9) and lacking femoral pores (versus 5���7). Other Thai species, including C. interdigitalis, C. intermedius, C. jarujini, C. oldhami, C. peguensis, C. pulchellus, and C. variegatus, were compared to the new species as above. Two described congeners occur in Cambodia: C. intermedius (Grismer et al. 2008; Smith 1935) and C. pseudoquadrivirgatus (Stuart et al. 2010). Cyrtodactylus bugiamapensis should occur in Cambodia because its habitat extends continuously from the type locality to the border between Vietnam and Cambodia (see also Nazarov et al. 2012). These three species were compared above to the new species. Etymology. The specific epithet puhuensis refers to the type locality, P�� Hu Nature Reserve, where the new species was discovered. Distribution and Habitat. Cyrtodactylus puhuensis sp. nov. is known only from the P�� Hu Nature Reserve, in northwestern Thanh H��a Province, northern Vietnam (Fig. 2). The single known specimen of this new gecko was found at night on a large tree (diameter about 1.5 m), at about 2.5 m above the forest floor. The habitat was evergreen forest intermixed with bamboo. Limestone was not observed in the area and the recorded elevation was 638 m., Published as part of Nguyen, Sang Ngoc, Yang, Jun-Xiao, Le, Thanh-Ngan Thi, Nguyen, Luan Thanh, Orlov, Nikolai L., Hoang, Chung Van, Nguyen, Truong Quang, Jin, Jie-Qiong, Rao, Ding-Qi, Hoang, Thao Ngoc, Che, Jing, Murphy, Robert W. & Zhang, Ya-Ping, 2014, DNA barcoding of Vietnamese bent-toed geckos (Squamata: Gekkonidae: Cyrtodactylus) and the description of a new species in Zootaxa 3784 (1), DOI: 10.11646/zootaxa.3784.1.2, http://zenodo.org/record/230341, {"references":["Nazarov, R., Poyarkov, N. A., Orlov, N. L., Phung, T. M., Nguyen, T. T., Hoang, D. M. & Ziegler, T. (2012) Two new cryptic species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from southern Vietnam. 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A., Sumontha, M., Ahmad, N., Bauer, A. M., Wangkulangkul, S., Grismer, J. L. & Pauwels, O. S. G. (2012) A phylogeny and taxonomy of the Thai-Malay Peninsula bent-toed geckos of the Cyrtodactylus pulchellus complex (Squamata: Gekkonidae): combined morphological and molecular analyses with descriptions of seven new species. Zootaxa, 3520, 1 - 55.","Sumontha, M., Panitvong, N. & Deein, G. (2010) Cyrtodactylus auribalteatus (Squamata: Gekkonidae), a new cave-dwelling gecko from Phitsanulok Province, Thailand. Zootaxa, 2370, 53 - 64.","Smith, M. A. (1923) Notes on reptiles and batrachians from Siam and Indo-China (No. 2). Journal of the Natural History Society of Siam, 6, 47 - 53.","Bauer, A. M., Sumontha, M. & Pauwels, O. S. G. (2003) Two new species of Cyrtodactylus (Reptilia: Squamata: Gekkonidae) from Thailand. Zootaxa, 376, 1 - 18.","Bauer, A. M., Kunya, K., Sumontha, M, Niyomwan, P., Nanitvong, P. & Pauwels, O. S. G. (2009) Cyrtodactylus erythrops (Squamata: Gekkonidae), a new cave-dwelling gecko from Mae Hong Son Province, Thailand. Zootaxa, 2124, 51 - 62.","Gray, J. E. (1831) A synopsis of the species of class Reptilia. In: Griffith, E. & Pidgeon, E. (Eds.), The animal kingdom arranged in conformity with its organisation by the Baron Cuvier with additional descriptions of all the species hither named, and of many before noticed. Whittaker, Treacher and Co., London, pp. 1 - 481 + 1 - 110.","Ulber, T. & Grossmann, W. (1991) Ein weiterer neuer Gecko aus Zentral-Thailand: Cyrtodactylus papilionoides sp. nov. (Reptilia: Sauria: Gekkonidae). Sauria, 13, 13 - 22.","Peters, W. (1871) Uber neue Reptilien aus Ostafrika und Sarawak (Borneo), vorzuglich aus der Sammlung des Hrn. Marquis J. Doria zu Genua. Monatsbericht der Koniglich Preußische Akademie der Wissenschaften zu Berlin, 1871, 566 - 581.","Boulenger, G. A. (1885) Catalogue of the Lizard in the British Museum (Natural History). Vol. 1. Trustees, London, 512 pp.","Pauwels, O. S. G., Sumontha, M., Latinne, A. & Grismer, L. L. (2013) Cyrtodactylus sanook (Squamata: Gekkonidae), a new cave-dwelling gecko from Chumphon Province, southern Thailand. Zootaxa, 3635 (3), 275 - 285. http: // dx. doi. org / 10.11646 / zootaxa. 3635.3.7","Bauer, A. M., Pauwels, O. S. G. & Chanhome, L. (2002) A new species of cave-dwelling Cyrtodactylus (Squamata: Gekkonidae) from Thailand. The Natural History Journal of Chulalongkorn University, 2, 19 - 29.","Chan-ard, T. & Makchai, S. (2011) A new insular species of Cyrtodactylus Gray, 1827 (Squamata, Gekkonidae), from the Surin Islands, Phang-nga Province, southern Thailand. The Thailand Natural History Museum Journal, 5, 7 - 15.","Pauwels, O. S. G., Bauer, A. M., Sumontha, M. & Chanhome, L. (2004) Cyrtodactylus thirakhupti (Squamata: Gekkonidae), a new cave-dwelling gecko from southern Thailand. Zootaxa, 772, 1 - 11.","Stuart, B. L., Rowley, J. J. L., Thy, N., Emmett, D. A. & Sitha, S. (2010) Significant new records of amphibians and reptiles from Virachey National Park, northeastern Cambodia. Cambodian Journal of Natural History, 2010, 38 - 47."]}

Li, Jia-Tang, Liu, Jun, Chen, Yue-Ying, Wu, Jia-Wei, Murphy, Robert W., Zhao, Er-Mi, Wang, Yue-Zhao, Zhang, Ya-Ping (2012): Molecular phylogeny of treefrogs in the Rhacophorus dugritei species complex (Anura: Rhacophoridae), with descriptions of two new species. Zoological Journal of the Linnean Society 165 (1): 143-162, DOI: 10.1111/j.1096-3642.2011.00790.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2011.00790.x

Rana megatympanum, new species (Previously referred to as species 7, ‘‘Large’’) Figures 12O, P, 13G, 14G, H HOLOTYPE: (ROM field no. 12999) ROM 39684, a gravid adult female from Khe Moi River, approximately 24 km west of Con Cuong village (by road), Con Cuong District, Nghe An Province, Vietnam (18°56̍30̎N, 104°48̍35̎E) found between 24 and 29 October 1994 by I.S. Darevsky, L.A. Lowcock, R.W. Murphy, and N.L. Orlov. The holotype had leg and liver tissue removed shortly after it was euthanised. PARATYPES: Eight females (ROM 39263, 39685–39691) collected with holotype between 24 and 29 October 1994 by I.S. Darevsky, L.A. Lowcock, R.W. Murphy, and N.L. Orlov. Four males (ROM 39237 –39240) from Na Hang Nature Reserve, Tuyen Quang Province, Vietnam (22°21̍54̎N, 105°25̍40̎E) found along waterfalls between 26 and 27 May 1996 by A. Lathrop and R.W. Murphy between 193 and 0200 hours. ROM 26398 – 26400 Con Cuong District, Nghe An Province, Vietnam (18°56̍30̎N, 104°48̍35̎E), collected 5 June 1995 by B. Hubley, A. Lathrop, R.W. Murphy, and N.L. Orlov. DIAGNOSIS: Rana megatympanum, a member of the subgenus Odorrana (sensu Fei et al., 1990), is characterized by a combination of the following attributes: (1) body dorsoventrally compressed; (2) SVL means of males 52 mm (48–55 mm), females 100 mm (93–105 mm); (3) vomerine teeth in rows oblique to choanae; (4) yellow lip­stripe present in males, absent or indistinct in females; (5) head broad, bluntly rounded in profile; (6) tympanum round, distinct, TMP:EYE in males enormous (1.20), greater than in females (0.51); (7) supratympanic fold weak; (8) dorsal skin shagreened, dorsolateral folds present only in males; (9) dorsum olive to brown sometimes with black spots; flanks marbled yellow and brown­gray; forelimbs and hindlimbs barred; (10) ventromedial callous pad on fingers II, III, and IV to proximal tubercle, fringes on fingers II, III, and IV; (11) disks on fingers and toes greatly enlarged (>2× base of phalanges); (12) feet fully webbed to toe disk, lateral fringes on I and V to terminal phalanges, webbing brown; (13) subarticular tubercles and an internal metatarsal tubercle distinct, conical; (14) terminal phalanges T­ shaped; (15) xiphisternum large, deeply notched posteriorly; (16) males with nuptial pads, paired gular pouches, pectoral spines absent; (17) eggs white. COMPARISONS: Rana megatympanum superficially resembles other Asian cascade ranids, including Huia nasica, Rana andersonii, R. archotaphus, R. bacboensis, R. banaorum, R. chalconota, R. chloronota, R. daorum, R. grahami, R. graminea, R. hainanensis, R. hejiangensis, R. hmongorum, R. hosii, R. morafkai, R. jingdongensis, R. junlianensis, R. kwangwuensis, R. leporipes, R. livida, R. margaretae, R. schmackeri, R. sinica, and R. tiannensis (table 12). It can be differentiated from all other Odorrana by the enormous sexual dimorphism in tympanum size (TMP:EYE 1.20 in males, 0.51 in females). Male R. megatympanum have a yellow lip­stripe and females have an indistinct or absent yellow lip­stripe, differentiating it from H. nasica, R. archotaphus, R. banaorum, R. chalconota, R. chloronota, R. daorum, R. graminea, R. hejiangensis, R. hosii, R. morafkai, R. leporipes, and R. livida (all with white lip­stripes); R. andersonii, R. bacboensis, R. hainanensis, R. jingdongensis, R. margaretae, and R. tiannensis have vertical lip­bars; R. junlianensis has a yellow lipstripe with brown lip­bars; R. schmackeri has no lip­stripe or vertical lip­bars. The broad, rounded snout differentiates R. megatympanum from H. nasica, R. andersonii, R. chalconota, and R. schmackeri (obtusely pointed) and from R. graminea and R. margaretae (depressed). Its gular pouches distinguish R. megatympanum from R. andersonii, R. chalconota, R. grahami, R. hainanensis, R. hmongorum, R. hosii, R. jingdongensis, R. junlianensis, R. kwangwuensis, and R. margaretae. Dorsolateral folds immediately differentiate male R. megatympanum from R. andersonii, R. bacboensis, R. chloronota, R. hainanensis, R. hejiangensis, R. morafkai, R. jingdongensis, R. junlianensis, R. kwangwuensis, R. livida, R. schmackeri, R. sinica, and R. tiannensis. The dorsolateral fold of male R. megatympanum differs from that of R. daorum, which is composed of minute white granules, R. hmongorum and R. grahami, whose dorsolateral pustules sometimes form a dorsolateral fold, and R. banaorum and R. chalconota, which have folds that are distinct and continuous, extending to the groin. The olive brown coloration with black spots of R. megatympanum differentiates it from H. nasica (olive­brown dorsum, lighter brown laterally) and from R. archotaphus, R. chalconota, R. chloronota, R. daorum, R. graminea, R. hejiangensis, R. hmongorum, R. hosii, R. jingdongensis, R. junlianensis, R. kwangwuensis, R. leporipes, R. margaretae, R. schmackeri, R. sinica (all with green). Rana megatympanum lacks an external metatarsal tubercle present in R. archotaphus and R. chalconota. Absence of ventral spines in male R. megatympanum separates them from R. andersonii, R. grahami, R. jingdongensis, R. junlianensis, R. margaretae, and R. schmackeri. The presence of white eggs differentiates R. megatympanum from H. nasica, R. andersonii, R. chalconota, R. grahami, R. junlianensis, R. margaretae, R. schmackeri (white eggs with melanic poles), and R. bacboensis (eggs completely melanic). Rana megatympanum differs from R. sinica by its distinct, uncovered tympanum (indistinct and covered with a layer of skin in R. sinica), its disk size (small in R. sinica), its relative finger lengths (I R. sinica, II R. megatympanum), and its T­ shaped distal phalanges (rounded in R. sinica). Rana megatympanum can be distinguished from R. leporipes by its webbing (only to the basal end of the distal phalanx in R. leporipes, to the disk in R. megatympanum), supratympanic fold (colored white in R. leporipes), and its T­ shaped distal phalanges (oblong, somewhat rounded in R. leopripes). Rana megatypmanum also differs from R. hejiangensis by its finger formula (II R. hejiangensis), and by its large disks (small for R. hejiangensis). Rana megatympanum closely resembles R. andersonii, R. hainanensis, R. jingdongensis, and R. tiannensis. In addition, R. megatympanum differs from R. andersonii in that its males are smaller (53–78 mm for males of R. andersonii), and its finger and toe­disks are relatively larger. Rana megatympanum differs from R. hainanensis in its smaller SVL (R. hainanensis males 49–62 mm, females 75–122 mm, R. megatympanum males 48–55 mm, females 93–105 mm) and its relative finger lengths (II R. hainanensis, II R. megatympanum). Rana megatympanum further differs from R. jingdongensis by its smaller males (R. jingdongensis SVL 62–81 mm) and skin (R. jingdongensis dorsum scattered with tubercles and large warts, lips and sides of heads with white spines, all absent in R. megatympanum). Rana megatympanum most closely resembles R. tiannensis, another large brown cascade ranid, but differs from it by having shagreened dorsal skin with small lateral granulations (dorsum of R. tiannensis is rough with large, prominent lateral granulations), and toe disks are smaller than those on fingers (the opposite condition of R. tiannensis). DESCRIPTION OF HOLOTYPE: ROM 39684, a gravid female, head width 77% of head length, length 50% of SVL; snout short, acutely rounded in dorsal view, bluntly rounded in profile, protruding beyond margin of lower jaw; eye very large, prominent, 73% of snout length; eyelid broader than interorbital distance. Top of head flat; canthus rostralis rounded; loreal region concave; lip flared just anterior to orbit; nostril about three­fourths distance from eye to tip of snout; supratympanic fold curving posteroventrally from posterior corner of eye to a level above the insertion of arm; tympanum round, distinctly visible, separated from eye by distance equal to TMP, 52% of EYE. Choanae ovoid; vomerine dentigerous processes prominent, oblique, posteromedial to choanae, each bearing numerous teeth. Tongue cordiform, distinctly notched posteriorly, free for approximately two­thirds its length. Forearms moderately robust; fingers moderately short, slender, hands 27% of SVL, relative lengths of fingers II 2× base of phalanges), relative pad size II IV k V, pad width (IV) 75% of pad length, each with ventral circummarginal grooves; subarticular tubercles prominent and conical; inner metatarsal tubercle ovoid, long; outer metatarsal tubercle absent. Xiphisternum large, deeply notched posteriorly. Skin on dorsum shagreened, becoming increasingly granular laterally; dorsolateral folds absent; small tubercles posteroventral to tympanum; prominent granules on flanks and around cloaca; cloacal opening unmodified, directed posteriorly, at upper level of thighs. COLOR IN LIFE (in preservative): Dorsum olive­brown, flanks yellow and brown­gray (gray to olive); lip­stripe absent (tympanum beige with dark brown center ring); loreal brown (black); iris gold; top one­third red, dorsal limbs brown with black banding (brown); posterior surface of thighs brown with black marbling (cloacal region black, thighs gray with white mottling); webbing marbled white on dark brown (brown on white); venter creamy white (creamy yellow with black mottling). SECONDARY SEXUAL CHARACTERS: The eggs of the holotype are creamy white and 2 mm in diameter. Adult females have SVL approximately twice that of males. Males have a yellow lip­stripe, and females either lack one or have an indistinctly yellow lip. Males also possess a weak dorsolateral fold, whereas females do not. Males have a larger tympanum than females, velvety nuptial pads extending across the thumb, paired gular pouches located at the angle of the jaw, and no pectoral spines. MEASUREMENTS OF HOLOTYPE (in mm): SVL 93.6; SNT 15.0; HDL 46.5; HDW 35.7; EYE 10.8; IOD 6.8; TMP 5.6; TEY 4.1; HND 25.7; FGR 19.6; FPL 3.0; FPW 2.6; TIB 65.0; FTL 49.7; TPL 3.2; TPW 2.4. VARIATION OF PARATYPES: Variation in all type material is given in table 15. MEASUREMENTS OF FEMALE PARATYPES (in mm, n = 10, ROM 26398 –26400, 39685– 39691): SVL 100.3 ± 4.2 (93.6–105.3); SNT 14.8 ± 0.8 (13.8–16.5); HDL 45.1 ± 3.3 (41.3–47.6); HDW 35.2 ± 0.7 (34.1–35.7); EYE 10.2 ± 0.7 (9.3–11.6); IOD 8.8 ± 1.0 (6.8–10.0); TMP 5.3 ± 0.4 (4.6–5.9); TEY 4.7 ± 0.3 (4.1–5.0); HND 25.2 ± 2.5 (20.6– 29.7); FGR 21.0 ± 1.0 (19.6–22.6); FPL 3.5 ± 0.6 (2.8–4.4); FPW 3.0 ± 0.5 (2.5–3.8); TIB 63.0 ± 3.3 (55.8–67.7); FTL 72.0 ± 12.1 (49.7–88.0) TPL 3.4 ± 1.0 (1.9–5.2); TPW 2.8 ± 0.4 (2.2–3.4). MEASUREMENTS OF MALE PARATYPES (in mm, n = 4, ROM 39237 –39240): SVL 52.3 ± 3.4 (48.6–55.2); SNT 8.7 ± 0.8 (8.3–9.6); HDL 28.0 ± 1.4 (24.6–27.1); HDW 18.5 ± 0.8 (18.0–19.1); EYE 4.1 ± 0.9 (3.17–4.6); IOD 5.0 ± 0.5 (4.7–5.6); TMP 4.7 ± 0.3 (4.3–5.1); TEY 1.7 ± 0.3 (1.5–2.0); HND 15.2 ± 0.4 (14.8–15.5); FGR 12.5 ± 0.4 (12.2–13.0); FPL 2.0 ± 0.5 (1.6–2.5); FPW 1.9 ± 0.4 (1.4–2.3); TIB 32.7 ± 0.4 (32.3– 33.1); FTL 38.3 ± 7.3 (29.9–42.8) TPL 1.8 ± 0.6 (1.3–2.5); TPW 1.6 ± 0.1 (1.5–1.6). ETYMOLOGY: The specific name is a noun in opposition, derived from the Latin prefix ‘‘mega’’ (meaning very large) and ‘‘tympanum’’, in reference to the relatively large tympanum of this species. DISTRIBUTION AND ECOLOGY: Rana megatympanum is known from northern and north­central Vietnam. It occurs in montane rivers that vary from shallow and slow moving to torrential and deep. It may be found on boulders and logs, both in and around the water as well as in the adjacent forest. Radiographs revealed that large invertebrates (both insects and crustaceans) form part of the diet. Vocalizations and tadpoles are unknown. REMARKS: The dorsolateral fold and very large tympanum of male R. megatympanum potentially make it Bourret’s (1942) ‘‘northern form’’ of R. chloronota (= R. graminea). However, R. graminea is bright green above (Boulenger, 1899) in contrast to the olivebrown dorsum of R. megatympanum.

Rana sinica Ahl, 1925 Figure 10 HOLOTYPE: ZMB 9785 an adult female from China. DIAGNOSIS: Rana sinica is characterized by a combination of the following attributes: (1) body dorsoventrally compressed; (2) SVL of single mature female is 66.6 mm, males unknown; (3) vomerine teeth in rows oblique to choanae; (4) lip��stripe absent; (5) head not broad, snout rounded in dorsal view; (6) tympanum round, indistinct, covered by a layer of skin, TMP:EYE 0.52; (7) supratympanic fold weak; (8) dorsal skin smooth flanks weakly granular, dorsolateral folds absent, venter smooth; (9) dorsum bronze��green with irregular indistinct spots and blue��gray flanks, legs with black bands; cloacal region marbled black with white; (10) median callous pad on finger III almost to proximal tubercle; (11) disks on fingers and toes slightly enlarged (Rana sinica superficially resembles other Asian cascade ranids, including Huia nasica, Rana andersonii, R. archotaphus, R. chalconota, R. chloronota, R. grahami, R. graminea, R. hainanensis, R. hejiangensis, R. hosii, R. jingdongensis, R. junlianensis, R. kwangwuensis, R. leporipes, R. livida, R. margaretae, R. schmackeri and R. tiannensis (table 12). Unlike the species listed above, R. sinica has a layer of skin overlying its tympanum and its terminal phalanges are slightly rounded (not T�� shaped; R. leporipes has oblong, somewhat rounded distal phalanges). The absence of a lip��stripe in R. sinica further differentiates it from H. nasica, R. archotaphus, R. chalconota, R. chloronota, R. grahami, R. graminea, R. hejiangensis, R. hosii, R. junlianensis, R. kwangwuensis, R. leporipes, and R. livida. Eggs of H. nasica, Rana andersonii, R. chalconota, R. grahami, R. junlianensis, R. margaretae, and R. schmackeri are white with melanic poles, whereas those of R. sinica are immaculate. The smooth green dorsum further distinguishes R. sinica from H. nasica, R. andersonii, R. jingdongensis, R. grahami, R. graminea, R. leporipes, R. schmackeri, and R. tiannensis. Huia nasica has an olive�� brown dorsum (R. sinica green). Rana hainanensis further differs from R. sinica in its size (mean female SVL R. hainanensis 103 mm) and relative lengths of fingers (II R. hainanensis, I R. sinica). Rana andersonii has a rough, olivebrown dorsum. Rana hejiangensis has smaller males (SVL 47 mm) and a different finger formula (II R. sinica. Rana julianensis and R. kwangwuensis differ from R. sinica in their relative finger formula (II Rana jingdongensis possesses vertical lip bands, large toe disks, and is fully webbed to all disks. Rana graminea differs from R. sinica in that its nares is closer to the tip of the snout than to the eye (halfway in R. sinica), and it has a different finger formula (II = I R. graminea). Rana margaretae further differs from R. sinica in having larger females (SVL 78���88 mm). Rana grahami has pustules on the dorsum and flanks and no digital disks (only slightly swollen tips). Rana schmackeri has a smooth, heavily spotted dorsum. Rana tiannensis has a rough, brown dorsum with large, prominent lateral granulations. Rana chalconota differs from R. sinica by having distinct dorsolateral folds, a pointed snout (vs. rounded), and an outer metatarsal tubercle. In R. archotaphus the outer metatarsal tubercle is present. Rana hosii differs from R. sinica by its dorsolateral folds and feeble tarsal folds (absent in R. sinica). Rana chloronota differs from R. sinica in being larger (SVL for adult females 80���100 mm for R. chloronota), having nostrils closer to the tip of the snout than the eye (halfway for R. sinica), and a different digital formula (II R. chloronota). Rana leporipes also differs from R. sinica by having a white supratympanic fold (not colored in R. sinica). Rana livida has a solid��colored dorsum with white spots on its sides (flanks a different color for R. sinica, lacking spots), and it lacks transverse bands on the arms and legs (present in R. sinica). DESCRIPTION OF HOLOTYPE: HDL greater than HDW (132%), HDW 26% of SVL, HDL 34% of SVL; snout short, rounded in dorsal view; protruding beyond margin of lower jaw, rounded in lateral view, EYE prominent, smaller than SNT, IOD broader than EYE and upper eyelid. Top of head flat; canthus rostralis rounded; loreal region steeply concave; nostril about one��half the distance from eye to tip of snout; supratympanic fold weak; tympanum round, indistinct, covered by layer of skin; TMP 52% of EYE in females. Choanae ovoid; vomerine dentigerous processes prominent, oblique, posteromedial to choanae, each bearing numerous teeth. Tongue cordiform, distinctly notched posteriorly, free for part of its posterior length. Forearm robust; relative lengths of fingers I R. sinica; all others are lost. This redescription is based on the original description by Ahl (1925) with amendments and additions based on our examination of the holotype. Ahl described the holotype as a male, but it is clearly a gravid female. As well, Ahl described a frog with large finger and toe pads. Although the condition of the specimen did not allow for accurate pad measurements to be made, the enlarged pads on the holotype are not as large as those on other species in the Rana chloronota complex. This species differs significantly from the other members of the Rana chloronota complex in preservative, but presumably has superficial resemblances in life. Ahl (1925) believed that R. sinica was closely related to Staurois natator., Published as part of BAIN, RAOUL H., LATHROP, AMY, MURPHY, ROBERT W., ORLOV, NIKOLAI L. & CUC, HO THU, 2003, Cryptic Species of a Cascade Frog from Southeast Asia: Taxonomic Revisions and Descriptions of Six New Species, pp. 1-60 in American Museum Novitates 3417 on pages 30-32, DOI: 10.1206/0003-0082(2003)4172.0.CO;2, http://zenodo.org/record/4734880, {"references":["Ahl, E. 1925 (1927). Ueber vernachlassigte Merkmale bei Froschen. Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin 1925 (1 - 10): 40 - 47."]}

Rana banaorum, new species (Previously referred to as species 6, ������Southern Big Eye������) Figures 12M, N, 13F, 14E, F HOLOTYPE: (ROM field no. 7145) ROM 39944, an adult female from Tram Lap, An Khe District, Gia Lai Province, Vietnam (14��26��39��N, 108��32��97��E), elevation ca. 900 m, collected on 15 June 1996 by R.H. Bain, A. Lathrop, R.W. Murphy, and N.L. Orlov. The holotype had leg and liver tissue removed shortly after it was euthanised. PARATYPES: Four males (ROM 39912 ��� 39913, 39915���39916) and eight females (ROM 39899 ���39901, 39928, 39929, 39931, 39936, and 39941) collected with holotype between 15 and 29 June 1996 by R.H. Bain, A. Lathrop, R.W. Murphy, and N.L. Orlov. ROM 39716 ���39720, 39920 ���39922, and 39924���39926 (males); 39942 (subadult) from the Cha River, Buon Loi, An Khe District, Gia Lai Province, Vietnam (elevation ca. 900 m), collected on 27 June 1996 by R.H. Bain and N.L. Orlov. ROM 39700, and 39702���39705 males from Krong Pa, An Khe District, Gia Lai Province, Vietnam (14��20��29�� N, 108��28��46��E, elevation 850 m), collected on 13���25 August 1997 by C.T. Ho, A. Lathrop, L.A. Lowcock, R.W. Murphy, and N.L. Orlov. ROM 25084 ���25086, 25100, 25102, and 25103 (males) collected from Buon Loi, An Khe District, Gia Lai Province, Vietnam (elevation ca. 700���750 m), 5 November 1993 by I.S. Darevsky and N.L. Orlov. DIAGNOSIS: Rana banaorum, a member of the subgenus Odorrana (sensu Fei et al., 1990), is characterized by a combination of the following attributes: (1) body dorsoventrally compressed; (2) SVL means of males 50 mm (42���55 mm), females 93 mm (83���99 mm); (3) vomerine teeth in rows oblique to choanae; (4) white lip��stripe extending across upper lip, terminating in glandule above insertion of arm; (5) head broad, bluntly rounded in profile; (6) tympanum round, distinct, very large, TMP:EYE in males (0.89) greater than females (0.75); (7) supratympanic fold weak; (8) dorsal skin shagreened, becoming granular laterally, with slight dorsolateral folds; (9) dorsum usually brown with yellow and black spots, sometimes green with black spots; legs brown, lightly banded; (10) median callous pad on fingers II, and III to proximal tubercle; (11) disks on fingers and toes greatly enlarged (>2�� base of phalanges); (12) feet fully webbed to disks, except medial side of IV, where it reaches disk as a fringe, lateral fringes on I and V to terminal phalanges, webbing brown; (13) subarticular tubercles and inner metatarsal tubercle distinct, conical; (14) terminal phalanges T�� shaped; (15) xiphisternum large, deeply notched posteriorly; (16) males with nuptial pads, paired gular pouches, pectoral spines absent; (17) eggs white. COMPARISONS: Among Asian cascade ranids, R. banaorum is most similar to sympatric R. morafkai and particularly R. chloronota but it can be anatomically distinguished from both by its dorsolateral folds (table 12). Rana banaorum can further be distinguished from R. morafkai by its larger males (SVL 42���55 mm, versus 39���45 mm). Some Rana banaorum males have indistinct, microscopic spinules on the dorsal surface of the leg extending to the feet, forming a ������saw��tooth������ formation along the lateral edge of toe V, which is present only to a slight degree or absent in R. chloronota. Its broad, rounded snout differs from the obtusely pointed snouts of H. nasica, R. andersonii, R. chalconota, and R. schmackeri and from the depressed snouts of R. graminea and R. margaretae. A white lip��stripe differentiates R. banaorum from R. andersonii, R. bacboensis, R. grahami, R. hainanensis, R. hmongorum, R. jingdongensis, R. junlianensis, R. margaretae, R. sinica, R. schmackeri, and R. tiannensis. The gular pouches of R. banaorum distinguish it from R. andersonii, R. chalconota, R. grahami, R. hainanensis, R. hmongorum, R. hosii, R. jingdongensis, R. junlianensis, R. kwangwuensis, and R. margaretae. The dorsolateral folds of R. banaorum distinguish it from H. nasica, R. andersonii, R. bacboensis, R. chloronota, R. hainanensis, R. hejiangensis, R. jingdongensis, R. junlianensis, R. kwangwuensis, R. morafkai, R. livida, R. schmackeri, R. sinica, and R. tiannensis; the dorsolateral folds of R. daorum are composed of distinct white granules, and dorsolateral pustules sometimes form folds on R. grahami and R. hmongorum. The absence of an outer metatarsal tubercle immediately differentiates R. banaorum from R. archotaphus and R. chalconota. The presence of white eggs differentiates R. banaorum from R. bacboensis (black eggs), Huia nasica, R. andersonii, R. chalconota, R. grahami, R. junlianensis, R. margaretae and R. schmackeri (white eggs with a dark melanic pole). The absence of spinules on the venter distinguishes R. banaorum from R. andersonii, R. grahami, R. jingdongensis, R. junlianensis, R. margaretae, and R. schmackeri. Rana banaorum further differs from Huia nasica by its larger females (83���99 mm vs. 67 mm in H. nasica). It is further differentiated from R. sinica by its uncovered, distinct tympanum (indistinct, covered by skin in R. sinica), large disks (small in R. sinica) and its relative finger lengths (I R. sinica, II R. banaorum) and T�� shaped distal phalanges (rounded in R. sinica). Rana leporipes further differs from R. banaorum with its white supratympanic fold (not colored in R. banaorum), webbing to distal phalanges (webbing to disks in R. banaorum), and T�� shaped distal phalanges (oblong, somewhat rounded in R. leporipes). DESCRIPTION OF HOLOTYPE: ROM 39944, an adult female, head width 78% head length, length 46% SVL; snout short, acutely rounded in dorsal view, bluntly rounded in profile, protruding beyond margin of lower jaw; eye large, prominent, 50% snout length; eyelid broader than interorbital distance. Top of head flat; canthus rostralis rounded; loreal region concave; lip flared just anterior to orbit; nostril about three��fourths distance from eye to tip of snout; supratympanic fold curving posteroventrally from posterior corner of eye to a level above the insertion of arm; tympanum very large, round, distinctly visible, separated from eye by distance equal to TMP, 80% of eye length. Choanae ovoid; vomerine dentigerous processes prominent, oblique, posteromedial to choanae, each bearing numerous teeth. Tongue cordiform, distinctly notched posteriorly, free for approximately two��thirds its length. Forearms moderately robust; fingers moderately short, slender, hands 23% SVL, relative finger lengths II 2�� base of phalanges), relative pad size II IV> V, pad length (IV) equals pad width; each pad with ventral circummarginal grooves; subarticular tubercles prominent and conical; inner metatarsal tubercle ovoid, long; outer metatarsal tubercle absent. Xiphisternum large, deeply notched posteriorly. Skin on dorsum shagreened, with small pustules on flanks; dorsolateral folds weak; small tubercles posteroventral to tympanum; prominent granules on flanks and around cloaca; cloacal opening unmodified, directed posteriorly, at upper level of thighs. COLOR IN LIFE (in preservative): Dark brown (olive��brown, beige), flanks gray with yellow spots (white spots); lip��stripe brown anteriorly and creamy white from level of the eye posteriorly (white); loreal dark brown (black); tympanum beige with dark brown central ring; iris golden brown; dorsal limbs grayish brown with black banding; cloacal region dark brown (black); webbing marbled white on dark brown (uniformly brown); venter creamy white, ventral surfaces of limbs creamy yellow with black mottling. SECONDARY SEXUAL CHARACTERS: The holotype is a gravid female with immaculate white eggs (2 mm in diameter). Gravid females have an SVL nearly twice that of males (mean SVL female, 93 mm; male 50 mm). Male TMP:EYE (0.89) is larger than females (0.75). Males have velvety nuptial pads on the thumb and paired gular pouches located at the angle of jaw. Pectoral spines are absent. MEASUREMENTS OF HOLOTYPE (in mm): SVL 98.0; SNT 13.5; HDL 45.4; HDW 35.6; EYE 6.8; IOD 7.4; TMP 5.4; TEY 4.7; HND 22.7; FGR 19.4; FPL 3.1; FPW 3.3; TIB 63.7; FTL 80.0; TPL 3.4; TPW 3.4. VARIATION OF PARATYPES: The skin on the dorsum varies from smooth to shagreened with small pustules on flanks. The lip��stripe varies from creamy yellow (white in alcohol) throughout its length to brown anteriorly and creamy white from the level of the eye posteriorly. The dorsal skin is variable from light green to dark brown (olive, brown, beige, or livid blue in alcohol) with or without large black spots. Variation in all type material is given in table 15. MEASUREMENTS OF FEMALE PARATYPES (in mm, n = 8, ROM 39899 ���39901, 39928, 39929, 39931, 39936, and 39941): SVL 92.7 �� 5.6 (83.4���98.7); SNT 13.8 �� 1.6 (12.4��� 16.8); HDL 43.8 �� 3.2 (37.5���48.8); HDW 30.6 �� 1.5 (26.7���33.4); EYE 6.1 �� 0.7 (4.7��� 6.9); IOD 8.2 �� 0.9 (6.8���10.1); TMP 4.5 �� 0.6 (3.5���5.6); TEY 4.5 �� 0.6 (3.5���5.2); HND 22.3 �� 2.8 (17.5���25.0); FGR 18.2 �� 2.8 (12.9���20.8); FPL 3.4 �� 0.4 (2.8���4.3); FPW 3.1 �� 0.2 (2.8���3.5); TIB 58.1 �� 3.2 (52.7���63.7); FTL 68.0 �� 7.0 (58.3���78.0) TPL 3.4 �� 0.6 (2.3���4.3); TPW 2.8 �� 0.4 (2.4���3.4). MEASUREMENTS OF MALE PARATYPES (in mm, n = 14, ROM 25084 ���25086, 25100, 25102, 25103, 39912, 39913, 39915, 39916, 39920���39922, 39924): SVL 50.5 �� 3.7 (42.5���54.6); SNT 7.4 �� 1.5 (3.1���8.7); HDL 26.1 �� 1.4 (24.6���28.1); HDW 17.4 �� 0.6 (17.8���18.3); EYE 4.0 �� 0.5 (3.1���5.1); IOD 3.8 �� 0.4 (3.4���4.7); TMP 3.5 �� 0.6 (2.0��� 4.7); TEY 1.7 �� 0.6 (0.9���3.6); HND 14.0 �� 1.2 (11.8���16.0); FGR 11.1 �� 1.1 (9.2���12.7); FPL 1.9 �� 0.3 (1.4���2.6); FPW 1.6 �� 0.2 (1.4���2.0); TIB 30.6 �� 4.6 (24.1���43.6); FTL 32.6 �� 6.8 (23.3���42.7); TPL 1.9 �� 0.4 (1.3��� 2.4); TPW 1.5 �� 0.3 (1.1���2.1). ETYMOLOGY: The specific name is a patronym for the Ba Na people, an ethnic group living on the Tay Nguyen Plateau (Central Highlands) of south��central Vietnam where this species occurs. DISTRIBUTION AND ECOLOGY: Rana banaorum is known only from the Tay Nguyen Plateau of the Central Highlands, Gia Lai Province, Vietnam. It inhabits forested montane river systems on or near rapids or waterfalls of primary and disturbed second growth. In May and June males have tight, leathery gular pouches, suggesting that they are not calling and that the breeding season occurs during some other time. REMARKS: Inger and Chanard (1997) and Inger et al. (1999) noted that specimens of Rana chloronota (as R. livida) from An Khe have more pronounced dorsolateral folds than elsewhere in Vietnam. These frogs do not fit Bourret���s (1942) interpretation of R. graminea as a ������northern variety������ of R. chloronota. These differences likely reflect the occurrence of three sympatric species of the R. chloronota complex from this region. Some male R. banaorum have indistinct, microscopic spinules on the dorsal surface of the leg extending to the feet, forming a ������saw��tooth������ formation along the lateral edge of toe (seen in FMNH specimens that are not part of the type series)., Published as part of BAIN, RAOUL H., LATHROP, AMY, MURPHY, ROBERT W., ORLOV, NIKOLAI L. & CUC, HO THU, 2003, Cryptic Species of a Cascade Frog from Southeast Asia: Taxonomic Revisions and Descriptions of Six New Species, pp. 1-60 in American Museum Novitates 3417 on pages 46-50, DOI: 10.1206/0003-0082(2003)4172.0.CO;2, http://zenodo.org/record/4734880, {"references":["Fei, L., C. - y. Ye, and Y. - z. Huang. 1990. Keys to Chinese Amphibians. Chongqing: Chongqing Branch, Science and Technology Materials Press, 364 pp. [in Chinese]","Inger, R. F., N. L. Orlov, and I. S. Darevsky. 1999. Frogs of Vietnam: a report on new collections. Fieldiana: Zoology (new series) 92: 1 - 46.","Bourret, R. 1942. Les batrachians de l'Indochine. Hanoi, Vietnam: Gouvernment General de L'Indochine, Hanoi, Vietnam, 547 pp."]}

Rana daorum, new species (Previously referred to as species 4, ‘‘Small’’) Figures 11E, F, 12G, H, 13C HOLOTYPE: (ROM field no. 19047) ROM 26381 an adult female from approximately 5 km NW of Sa Pa village, near O Qui Ho Pass, Lao Cai Province, Vietnam (22°22̍09̎N, 103°50̍14̎ E, elevation 1400 m) collected on 7 May 1995 along a waterfall beside the road by A. Lathrop and B. Hubley at approximately 1930 hours. PARATYPES: ROM 26382 –26397, all males, collected with the holotype, 7 May 1995 by A. Lathrop and B. Hubley at 1030 hours; ROM 38500 –38530, 38532–38540, 38542– 43, 38546–38561 collected between 30 April and 15 May 1999 in the vicinity of the type locality approximately 12 km northwest of Sa Pa Village near the O Qui Ho Pass (22°20̍58̎N, 103°46̍14̎E, elevation 1900 m) by R.O. de Sa´, C.T. Ho, A. Lathrop, T. Mason, R.W. Murphy, and N.L. Orlov. ROM 38547 is a subadult; ROM 38503, 38507, 38512, 38516, and 38530 are gravid females; ROM 38500, 38517, 38526, and 38538 are nongravid females; and the remaining specimens are males with distended gular pouches. DIAGNOSIS: Rana daorum, a member of the subgenus Odorrana (sensu Fei et al., 1990), is characterized by a combination of the following attributes: (1) body dorsoventrally compressed; (2) SVL means of males 36 mm (32–38 mm), females 55 mm (53–58 mm); (3) vomerine teeth absent; (4) lip­stripe white, extending across upper lip, terminating in a glandule above insertion of arm; (5) head not broad, snout rounded in dorsal view, rounded in profile; (6) tympanum round, distinct, TMP:EYE of females (0.45) greater than males (0.29); (7) supratympanic fold weak or absent; (8) dorsal skin smooth, granular near cloaca and tympanum, dorsolateral fold covered with small white granules, ventral skin smooth; (9) dorsum green occasionally with black spots, flanks brown with at least one large white glandular spot; fore­ and hindlimbs goldish brown, with mottling or indistinct transverse bands; (10) median callous pad on proximal two­thirds of fingers II and III; (11) disks on fingers and toes greatly enlarged (>2× base of phalanx), finger pads larger than toe pads; (12) feet fully webbed to disks, but as a fringe from distal subarticular tubercle of IV, lateral fringes on I and V to terminal phalanges, webbing mottled brown; (13) subarticular tubercles and inner metatarsal tubercle distinct, conical; (14) terminal phalanges T­ shaped; (15) xiphisternum narrow, forked posteriorly; (16) males with nuptial pads on thumb, paired gular pouches, pectoral spines absent; (17) eggs large, white. COMPARISONS: Though bearing a superficial resemblance to other cascade ranids of Southeast Asia, R. daorum is distinct (table 12) by the absence of vomerine teeth, females with larger TMP:EYE than males, and the presence of at least one large white spot on each flank. Its noticeably smaller SVL, dorsolateral folds formed by white granules, and solid, bright green dorsum immediately distinguishes it from H. nasica, R. andersonii, R. archotaphus, R. bacboensis, R. chloronota, R. grahami (whose dorsal pustules sometimes form a fold), R. graminea, R. hainanensis, R. hejiangensis, R. jingdongensis, R. junlianensis, R. kwangwuensis, R. livida, R. margaretae, R. schmackeri, R. sinica, and R. tiannensis. The entirely white eggs of R. daorum distinguish it from Huia nasica, Rana andersonii, R. chalconota, R. grahami, R. junlianensis, R. margaretae, R. schmackeri (all with white eggs with a melanic pole), and R. bacboensis (fully pigmented eggs). The presence of gular pouches in males differentiates R. daorum from R. andersonii, R. chalconota, R. grahami, R. hainanensis, R. hosii, R. jingdongensis, R. junlianensis, R. kwangwuensis, and R. margaretae. A distinct and uncovered tympanum also distinguishes R. daorum from R. sinica. Rana daorum can be further distinguished from R. livida, R. chloronota, R. maragaretae, and R. schmackeri by its finger formula (II R. daorum, I R. daorum closely resembles A. chunganenesis in size, the presence of white granular dorsolateral folds, and diurnal behavior, A. chunganensis is red­ brown, has vomerine teeth, and males have a TMP:EYE ratio twice that of R. daorum. DESCRIPTION OF THE HOLOTYPE: An adult female (ROM 26381), head width 74% of length, length 46% of SVL; snout short, rounded in dorsal view, rounded in profile, protruding beyond margin of lower jaw; eye very large, prominent, 72% of snout; eyelid broader than interorbital distance. Top of head flat; canthus rostralis rounded; loreal region vertical, concave; lip flared just anterior to orbit; nostril about three­fourths distance from eye to tip of snout; supratympanic fold indistinct, slight swelling above tympanum; tympanum round, distinctly visible, separated from eye by distance equal to that of TMP, 41% of EYE. Choanae ovoid; vomerine dentigerous processes absent. Tongue cordiform, distinctly notched posteriorly, free for approximately two­thirds its length. Forearms robust; fingers moderately short, slender, hand 28% of SVL, relative lengths of fingers I 2× base of phalanges), relative pad size II IV k V, pad width (IV) 83% of length, each pad with ventral circummarginal grooves; subarticular tubercles prominent and conical; inner metatarsal tubercle ovoid, long; outer metatarsal tubercle absent. Xiphisternum narrow, notched posteriorly. Skin on dorsum smooth, dorsolateral folds prominent in form of granules; small tubercles posteroventrally to tympanum, distinct granules on flanks and dorsum to cloaca; cloacal opening unmodified, directed posteriorly at upper level of thighs. COLOR IN LIFE (in preservative): Dorsum green (livid blue), flanks brown (brown) and green (gray), each a prominent white spot; granules of dorsolateral fold golden (white); lip­stripe white (silvery white) from nostril to above insertion of arm; tympanum dark brown; loreal region dark brown (black); dorsal surfaces of limbs mottled brown and yellow with indistinct dark brown (black) transverse bands; posterior surface of thighs yellow with brown (black) mottling; webbing marbled white (translucent) and dark brown; venter creamy white; iris golden, pupil outlined in a striking yellow and red border. SECONDARY SEXUAL CHARACTERS: Gravid females have immaculate white eggs. They are approximately 1.5 times larger than males. Males have a proportionally smaller tympanum than females (TMP:EYE for males 0.29, for females 0.45). The EYE:SNT is also greater in females (0.72) than it is in males (0.51). Males have velvety nuptial pads extending across thumb, and paired gular pouches located at the angles of the jaw. Pectoral spines are absent. MEASUREMENTS OF HOLOTYPE (in mm): SVL 55.7; SNT 7.8; HDL 25.4; HDW 18.8; EYE 5.6; IOD 5.6; TMP 2.3; TEY 2.4; HND 15.5; FGR 13.9; FPL 2.8; FPW 2.8; TIB 33.6; FTL 47.0; TPL 2.3; TPW 1.9. VARIATION OF PARATYPES: The loreal region on some specimens varies from dark brown to green. The large white spot on the flanks is sometimes accompanied by smaller ones. Flanks also have varying degrees of white mottling. The venter of some specimens has light mottling on the chest and chin. Variation in all type material is given in table 15. MEASUREMENTS OF FEMALE PARATYPES (in mm, n = 8, ROM 38500, 38503, 38507, 38512, 38516, 38517, 38526, 38530): SVL 55.0 ± 1.2 (53.3–57.6); SNT 7.3 ± 0.5 (6.8– 8.3); HDL 17.8 ± 1.6 (16.7–19.4); HDW 17.2 ± 0.6 (15.6–17.6); EYE 5.8 ± 0.4 (5.3– 6.4); IOD 10.0 ± 1.7 (10.0–11.3); TMP 2.5 ± 0.2 (2.3–3.0); FPW 2.3 ± 0.3 (1.8–2.8); TIB 34.3 ± 1.3 (32.7–36.4); TPW 2.3 ± 0.3 (1.8–2.8). MEASUREMENTS OF MALE PARATYPES (in mm, n = 7, ROM 26383, 26386, 26387, 26389, 26390, 26392, 26394): SVL 36.2 ± 1.2 (34.8–38.1); SNT 4.5 ± 0.4 (4.0–4.9); HDL 18.4 ± 1.7 (16.5–21.0); HDW 12.3 ± 0.6 (11.1–13.0); EYE 4.2 ± 0.6 (3.2–5.2); IOD 3 ± 0.4 (2.3–3.6); TMP 1.1 ± 0.2 (1.0– 1.9); TEY 0.6 ± 0.2 (0.2–1.0); HND 10.1 ± 0.6 (9.0–11.1); FGR 9.0 ± 0.3 (8.5–9.6); FPL 1.7 ± 0.3 (1.2–2.0); FPW 2.0 ± 0.2 (1.7–2.2); TIB 22.1 ± 1.3 (19.1–23.5); FTL 29.7 ± 4.4 (27.1–40.4); TPL 1.6 ± 0.2 (1.2– 1.7); TPW 1.4 ± 0.1 (1.2–1.6). ETYMOLOGY: The specific name is a patronym for the –Dao people (pronounced ‘‘zao’’) of northern Vietnam. DISTRIBUTION AND ECOLOGY: This species is known from the vicinity of Sa Pa village, Lao Cai Province in northern Vietnam. The photograph of a froglet of R. livida (in Karsen et al., 1998) also documents the occurrence of R. daorum in Hong Kong. The distinctive granular dorsolateral fold of R. daorum is clearly visible in the misidentified frog. In early May, male Vietnamese R. daorum are actively calling by 1000 hours on partially submerged rocks in cascades as well as in vegetation adjacent to the streams. Females, although not as common, can be found slightly farther away from streams in more dense vegetation. One male (ROM 26394) was found in amplexus with the holotype (fig. 6E). REMARKS: Rana daorum differs substantially from R. graminea (Boulenger, 1899; Bourret, 1942) despite having a dorsolateral fold. Its small, forked xiphisternum differs from the large, deeply notched element of other members in the Rana chloronota complex and the subgenus Odorrana (sensu Fei et al., 1990) and more closely resembles that of male Huia nasica (Yang, 1991b). Huia nasica shares range, habitat and morphological similarities with the Rana chloronota complex: greatly expanded finger and toe disks, tremendous sexual dimorphism in size, white eggs, paired gular pouches, and a high chirplike call (Boulenger, 1920; Pope, 1931; Bourret, 1942).

Rana hmongorum, new species (Previously referred to as species 3, ������Speckled������) Figures 14A, 12I, J, 13D HOLOTYPE: (ROM field no. 19055) ROM 26376 an adult female approximately 5 km NW of Sa Pa village, near O Qui Ho Pass, Lao Cai Province, Vietnam (22��20��09��N, 103��50��14��E, elevation 1400 m) collected 30 April 1995 by A. Lathrop and B. Hubley between 1900 and 2100 hours. The holotype had leg and liver tissue removed shortly after it was euthanised. PARATYPES: ROM 26370 ���26375, 26377��� 26380, 39235, and 39236 collected in the vicinity of the holotype between 9 and 12 May 1995 by A. Lathrop and B. Hubley. ROM 26380, 39235, and 39236 are males and the remainder females. Additional specimens from the vicinity of the type locality include AMNH A��161480 collected on 27 August 1997 by D.R. Frost and C.J. Raxworthy, and ROM 39867 ���39896 (22��20��58��N, 103��46��14��E, elevation 1900 m) collected between 30 April and 2 May 1999 by R.O. de Sa��, C.T. Ho, A. Lathrop, T. Mason, R.W. Murphy, and N.L. Orlov. Male specimens are ROM 39874 ���39879, 39888, and 39890���39894; juveniles and subadults ROM 39867, 39868, 39880, 39895, and 39896; all others are females. DIAGNOSIS: Rana hmongorum, a member of the subgenus Odorrana (sensu Fei et al., 1990), is characterized by a combination of the following attributes: (1) body dorsoventrally compressed; (2) SVL means of males 59 mm (54���65 mm), females 80 mm (74��� 87 mm); (3) vomerine teeth in rows posteromedial to choanae; (4) lip��stripe yellow, indistinct, extending across upper lip, terminating in a row of glandules; (5) head broad, bluntly rounded in profile; (6) tympanum round, distinct, TMP:EYE in males (0.43) greater than females (0.31); (7) supratympanic fold weak; (8) dorsal skin smooth with pustules becoming more pronounced laterally, pustules may form two dorsolateral lines resembling a fold, ventrum smooth; (9) dorsum and forearms green with black spots, flanks reddish brown with yellowish pustules, transverse bars on fore��� and hindlimbs; (10) median callous pad on fingers II and III to proximal tubercle; (11) disks on fingers and toes moderately expanded (Rana hmongorum superficially resembles other Asian cascade ranids, including Huia nasica, Rana andersonii, R. archotaphus, R. bacboensis, R. chalconota, R. chloronota, R. daorum, R. grahami, R. graminea, R. hainanensis, R. hejiangensis, R. hosii, R. jingdongensis, R. junlianensis, R. kwangwuensis, R. leporipes, R. livida, R. margaretae, R. schmackeri, R. sinica, and R. tiannensis (table 12). The white eggs of R. hmongorum distinguish it from H. nasica, Rana andersonii, R. chalconota, R. grahami, R. junlianensis, R. margaretae, R. schmackeri (all with white eggs with a melanic animal pole), and R. bacboensis (fully pigmented eggs). The dorsum and flanks with distinct pustules distinguishes R. hmongorum from all of the above species, except R. jingdongensis, and R. grahami; R. daorum has two distinct rows of granular dorsolateral folds. Absence of gular pouches further differentiates it from H. nasica, R. archotaphus, R. bacboensis, R. chloronota, R. daorum, R. graminea, R. junlianensis, R. kwangwuensis, R. schmackeri, and R. tiannensis. Rana hmongorum has an indistinct yellow lipstripe distinguishing it from all other cascade ranids above, except R. chalconota, R. jingdongensis, R. junlianensis, and R. grahami. The presence of webbing to the base of the toe pad distinguishes R. hmongorum from R. leporipes (webbing to distal phalanx) and R. archotaphus. Larger females and the absence of an outer metatarsal tubercle further distinguish R. hmongorum from R. archotaphus (female SVL 59���62 mm) and R. chalconota (SVL 46���59 mm). Absence of ventral spines in males differentiates R. hmongorum from R. andersonii, R. grahami, R. jingdongensis, R. junlianensis, R. margaretae, and R. schmackeri. Rana sinica is further differentiated from R. hmongorum by its indistinct, skin��covered tympanum, the relative length of its digits (I R. sinica, II R. hmongorum), and its rounded distal phalanges (T�� shaped in R. hmongorum). Huia nasica is different in that it has a smooth, olive��brown dorsum, a longer head and smaller size (67 mm vs.> 75 mm for females, 44���49 mm vs. 54���59 for males). The small male of R. hmongorum further distinguishes it from R. jingdongensis, R. junlianensis, R. andersonii, and R. grahami (SVL 62���81 mm for R. jingdongensis, 70��� 80 mm for R. andersonii, 66���84 mm for R. grahami). The relative length of fingers further distinguishes it from R. hainanensis (II R. hejiangensis (II IV> I> II, pad length (III) 85% of pad width; ventral circummarginal grooves on disks present; terminal phalanges T�� shaped; subarticular tubercles conical. Hindlimbs moderately robust; tibia length 57% of SVL; foot length 40% of SVL; relative toe lengths I IV k V, pad width (IV) 78% of pad length, ventral circummarginal grooves present; subarticular tubercles prominent and conical; inner metatarsal tubercle ovoid, long; outer metatarsal tubercle absent. Xiphisternum large, deeply notched posteriorly. Skin on dorsum smooth with large pustules, particularly on flanks, sacrum, and around cloaca; cloacal opening unmodified, directed posteriorly at upper level of thighs. COLOR IN LIFE (in preservative): Dorsum green (dark brown) and reddish brown (dark brown) with black spots. Sides reddish brown (gray) and yellow (cream), with some black spotting. Pustules on flanks yellowish white (creamy white), with black around the bases. Cloacal region dark brown (black). Lip��stripe creamy white (gray); dorsal surface of forearms with green (gray) patch; limbs mottled yellow (light brown) and brown (dark brown) with black transverse bands; anterior and posterior surfaces of thighs with brown (dark brown) marbling on yellow (cream); webbing uniformly brown; venter creamy white (creamy yellow). Iris greenish yellow or brown, pupil outlined with a yellow border. SECONDARY SEXUAL CHARACTERISTICS: Holotype is a gravid female with white eggs, 2 mm diameter. Adult female SVL 135% that of males. Males have slightly larger tympanum than do females, and velvety nuptial pads on thumb. Paired gular pouches and pectoral spines absent. Venter of males varies from immaculate creamy white to darkly mottled brown (gray in preservative). MEASUREMENTS OF HOLOTYPE (in mm): SVL 86.8; SNT 11.1; HDL 42.3; HDW 31.1; EYE 9.7; IOD 6.4; TMP 2.5; TEY 3.9; HND 22.3; FGR 16.5; FPL 2.3; FPW 2.7; TIB 49.2; FTL 35.0; TPL 3.0; TPW 2.3. VARIATION OF PARATYPES: Pustules can sometimes form dorsolateral lines, which superficially resemble folds. Pustules can be white. Variation in all type material is given in table 15. MEASUREMENTS OF FEMALE PARATYPES (in mm, n = 9; ROM 26370 ���26375, 26377��� 26379): SVL 80.2 �� 4.3 (74.3���86.8); SNT 11.0 �� 0.5 (10.1���11.7); HDL 38.8 �� 1.9 (37.0���41.0); HDW 29.8 �� 1.0 (29.2���31.3); EYE 9.3 �� 1.1 (7.6���10.8); IOD 7.0 �� 0.3 (6.4���7.3); TMP 2.9 �� 0.4 (2.5���3.3); TEY 3.0 �� 0.6 (2.1���3.9); HND 23.2 �� 1.3 (22.1��� 25.6); FGR 18.6 �� 1.1 (16.5���19.6); FPL 3.0 �� 0.4 (2.3���3.4); FPW 2.6 �� 0.4 (2.0���3.1); TIB 47.6 �� 2.3 (43.1���49.2); FTL 39.6 �� 5.0 (35.0���48.3) TPL 2.8 �� 0.6 (2.0���3.5); TPW 2.4 �� 0.5 (1.9���3.3). MEASUREMENTS OF MALE PARATYPES (in mm, n = 12; ROM 26380, 30876, 39235, 39236, 39874, 39875, 39877, 39879, 39888, 39891, 39892; AMNH A��161480): SVL 59.4 �� 3.8 (54.7���65.3); SNT 9.4 �� 0.9 (8.7��� 10.4); HDL 24.0 �� 3.0 (20.7���30.4); HDW 20.0 �� 1.2 (18.9���21.1); EYE 7.3 �� 0.5 (6.7��� 8.3); IOD 5.3 �� 0.7 (4.3���6.5); TMP 3.0 �� 0.4 (2.4���3.7); TEY 2.5 �� 0.3 (2.2���3.0); HND 17.2 �� 1.4 (14.1���19.4); FGR 13.3 �� 1.2 (12.8���14.8); FPL 2.5 �� 0.5 (1.6���3.6); FPW 2.1 �� 0.6 (1.0���2.4); TIB 38.6 �� 4.5 (33.4���51.2); FTL 47.2 �� 5.7 (30.4���52.1); TPL 2.3 �� 0.4 (1.7���2.9); TPW 1.7 �� 0.4 (0.9���2.2). ETYMOLOGY: The specific name is a patronym for the Hmong (pronounced ������huhmung������) people, an ethnic group in the northern montane regions of Vietnam. Their assistance made it possible for us to document the fauna of Hoang Lien Mountains. DISTRIBUTION AND ECOLOGY: This species is only known only from the vicinity of Mount Fan Si Pan at approximately 1900 m or above. Adults were found along the cascades of a waterfall and along the mossy slopes of a human��made culvert., Published as part of BAIN, RAOUL H., LATHROP, AMY, MURPHY, ROBERT W., ORLOV, NIKOLAI L. & CUC, HO THU, 2003, Cryptic Species of a Cascade Frog from Southeast Asia: Taxonomic Revisions and Descriptions of Six New Species, pp. 1-60 in American Museum Novitates 3417 on pages 40-44, DOI: 10.1206/0003-0082(2003)4172.0.CO;2, http://zenodo.org/record/4734880, {"references":["Fei, L., C. - y. Ye, and Y. - z. Huang. 1990. Keys to Chinese Amphibians. Chongqing: Chongqing Branch, Science and Technology Materials Press, 364 pp. [in Chinese]"]}

Rana livida (Blyth, 1856) Figures 6E, F, 7E, F, 9 Rana livida (Boulenger, 1887) NEOTYPE: BMNH 1889.3.25.48 an adult female from Thagata Juwa, Village on the hills southwest of Mt. Mooleyit, Dawna Mountains, Myanmar (400���500 m), by M.L. Fea in early 1887. DIAGNOSIS: Rana livida, a member of the subgenus Odorrana (sensu Fei et al. 1990), is characterized by a combination of the following attributes: (1) body dorsoventrally compressed; (2) SVL females 89.4, 97.1 mm; (3) vomerine teeth in rows oblique to choanae; (4) lip��stripe white, extending across upper lip, terminating in glandule above insertion of arm; (5) head broad, snout rounded in dorsal view, bluntly rounded in profile; (6) tympanum round, distinct, TMP:EYE 0.41; (7) supratympanic fold weak; (8) dorsal skin smooth to flanks, pustules dorsal to cloaca, dorsolateral folds absent, venter smooth; (9) dorsum uniform brown in preservative; forelimbs and hindlimbs lighter brown, without transverse bars; (10) median callous pad on finger III to proximal tubercle; (11) disks on fingers and toes enlarged (>2�� base of phalanges); (12) feet fully webbed to disks, weak lateral fringes on I and V to terminal phalanges; (13) subarticular tubercles and inner metatarsal tubercle distinct, conical; (14) terminal phalanges T�� shaped; (15) xiphisternum large, deeply notched posteriorly; (17) eggs white. COMPARISONS: Rana livida superficially resembles other Asian cascade ranids, including Huia nasica, Rana andersonii, R. archotaphus, R. chalconota, R. chloronota, R. grahami, R. graminea, R. hainanensis, R. hejiangensis, R. hosii, R. jingdongensis, R. junlianensis, R. kwangwuensis, R. leporipes, R. margaretae, R. schmackeri, R. sinica, and R. tiannensis (table 12). The smooth, uniformly colored dorsum further distinguishes R. livida from all of the above species. The unpigmented eggs of R. livida distinguish it from H. nasica, Rana andersonii, R. chalconota, R. grahami, R. junlianensis, R. margaretae, and R. schmackeri, which are white with melanic poles. Huia nasica differs from R. livida in its longer, more pointed head, and smaller females (SVL 67 mm). Rana hainanensis further differs from R. livida in relative lengths of fingers (II R. hainanensis, II R. livida). Rana hejiangensis differs from R. livida in its relative finger size (II Rana andersonii has a rough, olive��brown dorsum and small disks on fingers. Rana grahami has pustules on the dorsum and flanks (vs. smooth in R. livida) and slightly swollen digital tips. Rana junlianensis has brown lip bands. Rana kwangwuensis has small digital disks. Rana schmackeri has a smooth, heavily spotted dorsum. Rana tiannensis has a rough brown dorsum with large, prominent lateral granulations. Rana chalconota differs from R. livida by having coarsely shagreened dorsum, distinct dorsolateral folds, pointed snout, smaller SVL (females 46���59 mm), and an outer metatarsal tubercle. In R. archotaphus, webbing (toe IV) reaches base of distal subarticular tubercle (R. livida, webbing to base of disk), females are smaller (SVL 59���62 mm), and it has an outer metatarsal tubercle. Rana sinica differs from R. livida in lacking a lip��stripe, and in having smaller mature females (R. sinica holotype 66.6 mm), an indistinct, skin��covered tympanum (distinct and uncovered in R. livida), nares halfway between the eye and tip of the snout (nearer the snout in R. livida), and a different finger formula (I R. sinica, II R. livida). Rana hosii differs from R. livida by its obtusely pointed head (rounded in R. livida), dorsolateral folds, occasional bands on arms and legs, and feeble tarsal folds (dorsolateral and tarsal folds absent in R. livida). Rana chloronota differs from R. livida with its solid��colored dorsum and dark sides (uniform in R. livida), transverse bars on the limbs (absent in R. livida), smooth dorsum, flanks granular (slightly granulose on the posterior thighs in R. livida), and absence of white mottling on the flanks (present in R. livida). Rana leporipes differs from R. livida in having smaller digital disks, a white supratympanic fold, weak dorsolateral fold, and webbing that only reaches the distal phalanges (R. livida to base of the disk). Rana graminea further differs from R. livida with its nearly vertical (versus concave) loreal region and depressed head. DESCRIPTION: Head length greater than head width (103%), head width 35% of SVL, length 37% of SVL; snout short, protruding beyond margin of lower jaw, rounded in dorsal view, bluntly rounded in profile; eye large, prominent, 72% snout length; eyelid broader than interorbital distance. Top of head flat; canthus rostralis rounded; loreal region concave; lip flared just anterior to orbit; nostril about three��fourths distance from eye to tip of snout; supratympanic fold weak, swollen rim of tissue dorsal to tympanum; tympanum round, distinctly visible, 44% of EYE. Choanae ovoid; vomerine dentigerous processes prominent, oblique, posteromedial to choanae, each bearing numerous teeth. Tongue cordiform, distinctly notched posteriorly, free for approximately one��third its length. Forearm robust; fingers moderately short, slender; hands 29% of SVL, relative lengths of fingers II 2�� base of phalanges), relative pad size II IV> V; ventral circummarginal grooves present; subarticular tubercles prominent, conical; inner metatarsal tubercle ovoid, long; outer metatarsal tubercle absent. Xiphisternum large, deeply notched posteriorly. Skin on dorsum smooth to flanks; dorsolateral fold absent; small tubercles posteroventral to tympanum; granules dorsal cloaca; cloacal opening unmodified, directed posteriorly, at upper level of thighs. COLOR IN PRESERVATIVE: Dorsum uniform dark brown to flanks, some white spots and mottling on flank; prominent lip��stripe white; tympanum brown; posterior surface of thighs light brown with white spots, no transverse banding; venter creamy white; ventral side of limbs creamy white; webbing dark gray. SECONDARY SEX CHARACTERS: BMNH 1889.3.25.47, an adult female bears white eggs. No male specimens were studied. MEASUREMENTS OF NEOTYPE (in mm): SVL 89.4; SNT 14.2; HDL 32.8; HDW 31.6; EYE 10.2; IOD 8.0; TMP 4.5; TEY 2.8; HND 25.5; FPL 3.2; TIB 60.8; FTL 76.2; TPL 2.6. MEASUREMENTS OF REFFERED SPECIMEN (BMNH 1889.3.25.47): SVL 97.1; SNT 16.7; HDL 36.1; HDW 33.4; EYE 11.9; IOD 9.2; TMP 4.6; TEY 4.1; HND 28.8; FPL 3.3; TIB 63.8; FTL 75.6; TPL 2.7. DISTRIBUTION AND ECOLOGY: The range of R. livida is unclear. The type material is lost and the locality information has been poorly recorded. Much confusion exists in its taxonomy. The only confirmed locality is that of the neotype and maps do not show Thagata Juwa village, as indicated in by Feae. Mount Mooleyit is currently found on maps as Mulayit Taung (16��N, 98��30��E). The closest villages to Mulayit Taung are Kyeik��ywa, Daukkat��ywa, and Kyeik��don, to the west, and Mawkhi and Huthi, near the Thai border to the east. REMARKS: Male R. chloronota are substantially smaller than females and they posses paired subgular pouches and a larger tympanum. Presumably, male R. livida have the same attributes. Blyth (1852) appears to have included males in his original description; many of the specimens that Theobold assumed to be juveniles have whiter lip��stripes and a relatively large tympanum that is closer to the eye than in females. Although the color is not recorded in life, Blyth (1856) reported that R. livida is ������uniform duskyplumbeous above, probably dull olive green when alive.������ Boulenger (1920) recorded measurements of two R. livida females from Thagata, Tenasserim. It is not clear whether these two specimens are the same as those described above., Published as part of BAIN, RAOUL H., LATHROP, AMY, MURPHY, ROBERT W., ORLOV, NIKOLAI L. & CUC, HO THU, 2003, Cryptic Species of a Cascade Frog from Southeast Asia: Taxonomic Revisions and Descriptions of Six New Species, pp. 1-60 in American Museum Novitates 3417 on pages 28-30, DOI: 10.1206/0003-0082(2003)4172.0.CO;2, http://zenodo.org/record/4734880, {"references":["Blyth, E. 1856. Report for the October meeting, 1855. Journal of the Asiatic Society of Bengal 25 (7): 711 - 723.","Boulenger, G. A. 1887. An account of the reptiles and batrachians obtained in Tenasserim by M. L. Feae, of the Genoa Civic Museum. Annali dell Museo Civico di Storia Naturale di Genova, ser. 2, 4: 474 - 486, 3 pls.","Fei, L., C. - y. Ye, and Y. - z. Huang. 1990. Keys to Chinese Amphibians. Chongqing: Chongqing Branch, Science and Technology Materials Press, 364 pp. [in Chinese]","Blyth, E. 1852. Proceeding of the Society for April. Journal of the Asiatic Society of Bengal 21: 351 - 356.","Boulenger, G. A. 1920. A monograph of the south Asian, Papuan, Melanesian, and Australasian frogs of the genus Rana. Records of the Indian Museum 20: 1 - 126."]}

KEY TO VIETNAMESE SPECIES IN THE RANA CHLORONOTA COMPLEX The Rana chloronota complex in Vietnam includes species that are dorsoventrally compressed with long legs. Snout���vent length of females is approximately twice that of males (table 14). The dorsum may be green, brown, or a combination thereof; dorsum usually smooth, sometimes shagreened, often becoming more rugose laterally. Tympanum is distinct. Venter is entirely smooth. Legs with transverse bands or mottled with indistinct bands, posterior surface of thighs marbled yellow and black. An outer metatarsal tubercle is absent. Disks on fingers and toes well dilated, with ventral circummarginal grooves. Webbing is complete or nearly so, extending to most distal tubercle or to base of toe pad (as a fringe in some species). Males have velvety nuptial pads. VARIABLE CHARACTERS: Lip��stripe is present in most species and is usually white. Vomerine teeth, if present, are in rows oblique to the internal choanae. Males usually have paired gular pouches. Eggs completely pigmented or unpigmented. HABITAT: Ubiquitous throughout highland waterways of Southeast Asia. Found in and around water, including the forest floor and canopy of surrounding forests. Commonly found perched on rocks beside or among the cascades. 1. Dorsolateral fold distinct, composed of small gold (white in preservative) granules; noticeably smaller than other cascade ranids (males 35 mm, females 55 mm), large white lateral spot, and vomerine teeth absent.............................. R. daorum ��� Dorsolateral fold not as above or granules absent.............................. 2 2. Indistinct glandular dorsolateral folds present................................. 3 ��� Dorsolateral folds absent............... 4 3A. Lip��stripe indistinct yellow or absent in females, yellow in males; male tympanum 120% of eye length, 51% for females; skin completely shagreened, brown, and snout��� vent length notably large (males 52 mm, females 100 mm)... R. megatympanum 3B. White lip��stripe present in both males and females; dorsum green to olive, shagreened or incompletely smooth, flank gray; male tympanum 89% eye, females 75%; SVL males 50 mm, females 93 mm.......................... R. banaorum 3C. Dorsum bright green with or without black spots, completely smooth; flank brownish; male tympanum 77% eye, females 56%; SVL males 46 mm, females 94 mm........................... R. graminea 4. Dorsum completely smooth; dorsum bright green, vomerine teeth present, white lipstripe present, eggs immaculate, and SVL females 93 mm, males 46 mm............................. R. chloronota ��� Dorsum in part smooth and shagreened, or with heavy granulations............ 5 5. Dorsum with heavy granulations; skin heavily granulated laterally and on pelvis, dorsum bright green with black spots, males without gular pouches, SVL females 80 mm, males 60 mm...................................... R. hmongorum ��� Dorsum shagreened or in part smooth... 6 6. Black vertical bar on lip; dorsum shagreened, brown with some black spots, lip bands black, extend vertically across mandible, (white lip��stripe absent), webbing marbled white on dark brown to toes, eggs pigmented black, SVL 95 females, 55 mm males)................ R. bacboensis ��� No black vertical bar on lip; dorsum shagreened or incompletely smooth, color variable green to brown in daylight, changing to brown at night, white lip��stripe present, webbing brown, eggs immaculate white, SVL females 88 mm, males 43 mm........................... R. morafkai, Published as part of BAIN, RAOUL H., LATHROP, AMY, MURPHY, ROBERT W., ORLOV, NIKOLAI L. & CUC, HO THU, 2003, Cryptic Species of a Cascade Frog from Southeast Asia: Taxonomic Revisions and Descriptions of Six New Species, pp. 1-60 in American Museum Novitates 3417 on pages 53-54, DOI: 10.1206/0003-0082(2003)4172.0.CO;2, http://zenodo.org/record/4734880

Rana leporipes Werner, 1930 HOLOTYPE: Two figures (Werner, 1930: 49, pl. IV) of Mell no. 15660, an adult female from Lung Tao Shan, North Kwangtung, China (700 m a.s.l.) collected on 4 July 1919 by R. Mell. The voucher specimens of the type series have been lost and the photographs of the holotype become the iconotype. DIAGNOSIS (from text and plates of original description): Rana leporipes is characterized by a combination of the following attributes: (1) body dorsoventrally compressed; (2) SVL reported for females between 52 and 102 mm; (3) vomerine teeth in rows oblique to choanae; (4) lip��stripe white; (6) tympanum very distinct, TMP:EYE is 0.5; (7) supratympanic fold milky white; (8) dorsal skin smooth, flanks weakly granular, dorsolateral folds slightly distinguishable, venter smooth; (9) dorsum dark green, flanks stony gray with white marbling, legs not banded; (11) disks on fingers and toes only slightly enlarged (Rana leporipes superficially resembles other Asian cascade ranids, including Huia nasica, Rana andersonii, R. archotaphus, R. chalconota, R. chloronota, R. grahami, R. graminea, R. hainanensis, R. hejiangensis, R. hosii, R. jingdongensis, R. junlianensis, R. kwangwuensis, R. livida, R. margaretae, R. schmackeri, R. sinica and R. tiannensis (table 12). Rana leporipes is unique among all of the above species in having a white supratympanic fold, webbing that only reaches the distal phalanx (not the disk), and oblong, somewhat rounded distal phalanges (T�� shaped in others, unknown for R. hejiangensis, R. junlianensis, and R. kwangwuensis). The absence of banding on the legs distinguishes R. leporipes from all species listed here, except R. livida (present or absent in R. chalconota and R. hosii). The presence of a dorsolateral fold also distinguishes it from R. andersonii, R. chloronota, R. hainanensis, R. hejiangensis, R. jingdongensis, R. junlianensis, R. kwangwuensis, R. livida, R. margaretae, R. schmackeri, R. sinica, and R. tiannensis. Whereas Huia nasica has an olive��brown dorsum, R. leporipes is dark green. Rana hainanensis further differs from R. leporipes in its size (SVL R. hainanensis 103 mm) and relative lengths of fingers (II R. hainanensis, I = II for R. leporipes). Rana junlianensis has brown lip bands. Rana andersonii has a rough, olive��brown dorsum. Rana grahami has pustules on the dorsum and flanks and no digital disks. Rana schmackeri has a smooth, heavily spotted dorsum. Rana tiannensis has a rough, brown dorsum with large, prominent lateral granulations. Both Rana chalconota and R. archotaphus possess an outer metatarsal tubercle, but R. leporipes does not. Rana hosii can be distinguished from R. leporipes by its feeble tarsal folds. Rana chloronota differs from R. leporipes in that it is larger (R. chloronota female SVL 80���100 mm) and it has a different digital formula (II R. chloronota). Dorsum of R. livida is uniform (flanks a different color for R. leporipes). REMARKS: The original description of R. leporipes reported extensive variation in snout���vent length (52���102 mm) and lacked any description of secondary sex characters. The single male specimen of R. leporipes is significantly larger than males of R. chloronota (93 mm vs. a maximum of 53 mm). However, it is unclear how the male was sexed, as nuptial excrescences and gular pouches are not mentioned. Bourret (who did not indicate whether he had seen the type series) thought the male was an incorrectly identified female and that Werner���s (1930) frogs were all females spanning a range of varying degrees of maturity. We are also skeptical about any males, although this cannot be confirmed, as the type series has been lost., Published as part of BAIN, RAOUL H., LATHROP, AMY, MURPHY, ROBERT W., ORLOV, NIKOLAI L. & CUC, HO THU, 2003, Cryptic Species of a Cascade Frog from Southeast Asia: Taxonomic Revisions and Descriptions of Six New Species, pp. 1-60 in American Museum Novitates 3417 on pages 27-28, DOI: 10.1206/0003-0082(2003)4172.0.CO;2, http://zenodo.org/record/4734880, {"references":["Werner, F. 1930. Rana leporpies, a new species of frog from south China. Lignan Science Journal 9 (1, 2): 45 - 47, pl. 49."]}