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An integrative analysis recovered a new species of the Hemiphyllodactylus typus group from a karst formation in Lung Cu Commune, Dong Van District, Ha Giang Province, northeastern Vietnam. Hemiphyllodactylus lungcuensis sp. nov. is embedded within clade 6 of the typus group, bearing an uncorrected pairwise sequence divergence of 4.6–20.2% from all other species based on a 1,038 base pair segment of the mitochondrial NADH dehydrogenase subunit 2 gene (ND2). It is diagnosable from other species in clade 6 by statistically significant mean differences in normalized morphometric, meristic, and categorical characters. A multiple factor analysis using the three aforementioned character types recovered its unique, non-overlapping placement in morphospace as statistically significantly different from that of all other species in clade 6. The description of this new Hemiphyllodactylus species contributes to a growing body of literature underscoring the high degree of herpetological diversity and endemism in karst landscapes in Vietnam as well as in the genus Hemiphyllodactylus.

Cyrtodactylus arndti sp. nov. urn:lsid:zoobank.org:act: 83F60753-8E51-49D9-97E7-BD870AC772F3 Figs 6–9; Table 3 Diagnosis The new species of Cyrtodactylus is distinguished from remaining congeners of the C. irregularis species group by a combination of the following characters: SVL: 73.4–80.9 mm; dorsal pattern with 6 or 7 irregularly shaped bands; moderately broad nuchal band; original tail with irregular transverse bands; subcaudals transversely enlarged; 5–11 enlarged femoral scales; males with 0–2 pitted femoral pores, those absence in females; males with 6 precloacal pores, females with 6 pitted precloacal pores, pore-bearing scales arranged in a single ˄-shaped series; ventral scales 26–38; dorsal tubercles in 17–20 irregular longitudinal rows; precloacal groove absent; supralabials 8–13; infralabials 8–12; number of subdigital lamellae on fourth finger 15–20 and on fourth toe 17–22. Etymology We name this species in honor of our colleague, Prof. Dr. Hartmut Arndt, Institute of Zoology, University of Cologne, Germany, in recognition of his support for biodiversity research in Vietnam. As common names, we suggest Arndt’s Bent-toed Gecko (English) and Thằn lằn ngón arndt (Vietnamese). Type material (Figs 6–9) Holotype VIETNAM • ♂; Binh Dinh Province, Van Canh District, near Hiep Ha Village; 13°39.858′ N, 108°53.355′ E; 270 m a.s.l.; 13 Aug. 2016; D.T. Do and T.V. Nguyen; Field No. BD.2016.141; IEBR R.4930. Paratypes VIETNAM • 1 ♂; Binh Dinh Province, Quy Nhon District, Quy Nhon City; 13°41.718′ N, 109°10.277′ E; 140 m a.s.l; 8 Aug. 2016; D.T. Do and T.V. Nguyen; Field No. BD.2016.1; IEBR R.5219 • 2 ♂♂, 1 ♀; Binh Dinh Province, Van Canh District, near Dak Dum Village; 13°38.365′ N, 108°57.863′ E; 150 m a.s.l.; 11 Aug. 2016; D.T. Do and T.V. Nguyen; Field No. BD.2016.86, BD2016.87, BD.2016.88; IEBR R.4931 to IEBR.4933 • 1 ♀; Binh Dinh Province, Van Canh District, near Hiep Ha Village; 13°39.858′ N, 108°53.355′ E; 270 m a.s.l.; D.T. Do and T.V. Nguyen; Field No. BD.2016.86; IEBR R.5077 • 2 ♂♂; same collection data as for preceding; Field No. BD.2016.142, BD.2016.143; ZFMK 103910, ZFMK 103911. Description of holotype Adult male; snout-vent length 74.4 mm; tail regenerated, 98.15 mm in length; body slender, elongate (AG/SVL ratio 0.39); head distinct from neck, elongate, depressed (HL/SVL 0.29, HW/HL 0.67, HH/ HL 0.37); loreal region concave; snout long, blunt in dorsal profile (SE/HL 0.37), longer than diameter of orbit (OD/SE 0.61); scales on snout small, round or oval, granular, lager than scales on occiput; orbit large (OD/HL 0.22); pupils vertical; ear opening small, oval (ED/HL 0.06); rostral almost twice as wide as high with an inverse Y-shapted structure, surrounded by first supralabial, naris, nasorostral on each side, and internasal; nostril opening small and oval, surrounding by rostral, nasorostral, 2 supranasals and one postnasal; mental scale triangular, wider than high (ML/MW 0.74); two enlarged, triangular postmentals; supralabials 12/12; infralabials 11/13. Dorsal scales granular, dorsal tubercles round, keeled, conical, in 20 irregular rows at midbody; tubercles on occiput small; each tubercle surrounded by 9 or 10 granular scales; ventral scales smooth, round, midventral scales approximately 3–4 times as large as dorsal scales, slightly imbricate laterally, in 37 longitudinal rows at midbody between ventrolateral folds; precloacal groove absent; enlarged femoral scales 7 or 8 on each thigh, about twice the size of surrounding scales; enlarged precloacal scales 17, arranged in a rhombus; femoral pores absent; precloacal pores 6, arranged in ˄-shaped series. Fore and hindlimbs moderately slender (ForeaL/SVL 0.15 mm, CrusL/SVL 0.16); forelimbs dorsally covered by several slightly enlarged tubercles; dorsal surface of hindlimbs bearing well-developed tubercles; two postcloacal tubercles on each side on the hemipenal swellings; phalanges without webbing; each claw sheathed by two scales, the ventral sheath larger than the upper scale; number of subdigital lamellae on first finger 12/13, on first toe 13/13, on fourth finger 18/18, on fourth toe 22/22. Coloration in preservative Dorsal surface of head, body and limbs light-brown with some dark-brown pattern, without light bordering; occiput marbled with small, irregular dark-brown banded; rostral, mental, first three supralabials and first infralabials greyish brown, remaining infralabials light beige, some with greyish brown speckles or frames; nuchal band discontinuous, consisting of two stripes extending from the orbits to the neck, ending by a dark blotch on each side and a third blotch medially; dorsum with 7 irregular bands, the first two interrupted; dorsolateral region covered with round or elongate dark-brown blotches; tail with 6 dark-brown bands, fade ventrally, some small dark-brown spots arranged in a line along the lateral side of tail, tail tip dark-brown; dorsal surface of limbs with 6 or 7 irregular dark-brown bands; phalanges brown with beige knuckles; dorsal tubercles white or dark-brown depending on position; tubercles on dorsal surface of limbs and tail light-brown; venter greyish brown. Sexual dimorphism and variation The female (IEBR R.4933) differs from the males by the absence of hemipenal swellings. All male specimens have 6 precloacal pores but the females has 6 pitted scales only. Three males (IEBR R.4930, ZFMK 103910, ZFMK 103911) and the female lack femoral pores (IEBR R.5077, IEBR R.4933). For further morphological characters see Table 3. Comparisons The new species can be distinguished from all other member of Cyrtodactylus irregularis group from Vietnam by morphological characteristics (see Table 4). Cyrtodactylus arndti sp. nov. differs from C. badenensis by having the presence of enlarged femoral scales (5–11 vs absent in C. badenensis), the presence of precloacal pores in males (6 vs absent in C. badenensis), the presence of pitted precloacal pores in females (6 vs absent in C. badenensis), and different dorsal pattern (irregular bands vs banded in C. badenensis); differs from C. bidoupimontis by having the presence of pitted precloacal pores in females (6 vs absent in C. bidoupimontis), having moderately broad nuchal band, continuous or discontinuous band (vs well developed, widened posteriorly in C. bidoupimontis), different dorsal pattern (6 or 7 irregular transverse bands, colouration dark-brown on light-brown background vs 4–5 dark irregular transverse dorsal bands, usually with light borders), and the presence of transversely enlarged subcaudals vs absent in C. bidoupimontis; differs from C. bugiamapensis by having fewer precloacal pores in males (6 vs 7–11 in C. bugiamapensis), different dorsal color pattern (irregular, dark-brown transverse bands vs unclear transversal bands formed by irregular roundish to oblong, dark brown spots in C. bugiamapensis), moderately broad nuchal band, continuous or discontinuous bands (vs dark nuchal band, which can be medially divided, narrow, U-shape in C. bugiamapensis), and the presence of transversely enlarged subcaudals vs absent in C. bugiamapensis; differs from C. buchardi by having a lagger size (SVL of 73.4–80.9 vs 60–65 in C. buchardi), the presence of enlarged femoral scales (5–11 vs absent in C. buchardi), more subdigital lamellae on first finger (15–20 vs 14 in C. buchardi), more subdigital lamellae on first toe (17–22 vs 12 in C. buchardi) and the presence of transversely enlarged subcaudals vs absent in C. buchardi); differs from C. caovansungi by having a smaller size (SVL of 73.4–80.9 vs 90.4–94.0 mm), fewer femoral pores (0–2 vs 6 in C. caovansungi), fewer precloacal pores in males (6 vs 9 in C. caovansungi), the presence of pitted precloacal pores in females (6 vs absent in C. caovansungi), and fewer subdigital lamellae under the fourth toe (17–22 vs 23–25 in C. caovansungi); differs from C. cattienensis by having a larger size (73.4–80.9 mm vs 43.5–69.0 mm in C. cattienensis), the presence of pitted precloacal pores in females (6 vs absent in C. cattienensis), and the presence of transversely enlarged subcaudals (vs absent in C. cattienensis); differs from C. chungi by having a larger size (73.4–80.9 mm vs 66.6– 68.5 mm in C. chungi), fewer precloacal pores in males (6 vs 7 in C. chungi), fewer enlarged precloacal scales (17–23 vs 41–45 in C. chungi), and the presence of transversely enlarged subcaudals (vs absent in C. chungi); differs from C. cryptus by having fewer ventral scale rows (26–38 vs 47–50 in C. cryptus), the presence of enlarged femoral scales (5–11 vs absent in C. cryptus), fewer precloacal pores in males (6 vs 9–11 in C. cryptus), the presence of pitted precloacal pores in females (6 vs absent in C. cryptus), and the presence of transversely enlarged subcaudals (vs absent in C. cryptus); differs from C. cucdongensis by having a larger size (SVL 73.4–80.9 mm vs 55.8–65.9 mm in C. cucdongensis), fewer ventral scale rows (26–38 vs 41–44 in C. cucdongensis), more subdigital lamellae under the first toe (11–14 vs 8–11 in C. cucdongensis), and the presence of transversely enlarged subcaudals (vs absent in C. cucdongensis); differs from C. culaochamensis by having fewer ventral scale rows (26–38 vs 45– 50 in C. culaochamensis), the presence of enlarged femoral scales (5–11 vs absent in C. culaochamensis), fewer precloacal pores in males (6 vs 7–8 in C. culaochamensis), and the presence of pitted precloacal pores in females (6 vs absent in C. culaochamensis); differs from C. dati by having fewer ventral scale rows (26–38 vs 42–48 in C. dati), fewer femoral pores (0–2 vs 3–4 on each side in C. dati), different dorsal color pattern (irregular bands vs blotches in C. dati), and the presence of transversely enlarged subcaudals (vs absent in C. dati); differs from C. gialaiensis by having a larger size (73.4–80.9 mm vs 50.1–62.8mm in C. gialaiensis), the presence of enlarged femoral scales (5–11 vs absent in C. gialaiensis), fewer precloacal pores in males (6 vs 9–10 in C. gialaiensis), and more subdigital lamellae under the fourth finger (15–20 vs 14–15 in C. gialaiensis); differs from C. huynhi by having fewer ventral scale rows (26–38 vs 43–46 in C. huynhi), more enlarged femoral scales (5–11 vs 3–5 in C. huynhi), fewer precloacal pores in males (6 vs 7–9 in C. huynhi), and the presence of transversely enlarged subcaudals (vs absent in C. huynhi); differs from C. irregularis by having the presence of precloacal pores in males (1–2 vs absent in C. irregularis), different dorsal color pattern (irregular bands vs blotched in C. irregularis), and the presence of transversely enlarged subcaudals (vs absent in C. irregularis); differs from C. kingsadai by having a smaller size (SVL 73.4–80.9 mm vs 83.0–94.0 mm in C. kingsadai), fewer ventral scale rows (26–38 vs 39–46 in C. kingsadai), and fewer precloacal pores in males (6 vs 7–9 in C. kingsadai); differs from C. orlovi by having a larger size in males (SVL 73.4–80.9 mm vs 61.0– 68.2 mm in C. orlovi), the presence of pitted precloacal pores in females (6 vs absent in C. orlovi), and the presence of transversely enlarged subcaudals(vs absent in C. orlovi); differs from C.phnomchiensis by having fewer ventral scale rows (26–38 vs 45–54 in C. phnomchiensis), more precloacal pores in males (6 vs 4–5 in C. phnomchiensis), fewer AG/SVL ratio (0.34–0.43 vs 0.45–0.48 in C. phnomchiensis), and the presence of transversely enlarged subcaudals (vs absent in C. phnomchiensis); differs from C. phumyensis by having a larger size (SVL 73.4–80.9 mm vs 63.6–66.8 mm in C. phumyensis), and the presence of transversely enlarged subcaudals (vs absent in C. phumyensis); differs from C. phuocbinhensis by having a larger size (SVL 73.4–80.9 mm vs 46.0– 60.4 mm in C. phuocbinhensis), fewer ventral scale rows (26–38 vs 43–47 in C. phuocbinhensis), fewer precloacal pores in males (6 vs 7 in C. phuocbinhensis), the presence of pitted precloacal pores in females (6 vs absent in C. phuocbinhensis); differs from C. pseudoquadrivirgatus by having fewer ventral scale rows (26–38 vs 41–57 in C.pseudoquadrivirgatus), the presence of enlarged femoral scales (5–11 vs absent in C. pseudoquadrivirgatus), different dorsal color pattern (irregular bands vs blotched in C. pseudoquadrivirgatus), and the presence of transversely enlarged subcaudals (vs absent in C. pseudoquadrivirgatus); differs from C. raglai by having a smaller size (SVL 73.4–80.9 mm vs 95–111.7 mm in C. raglai), more precloacal pores in males (6 vs 5 in C. raglai) and the presence of pitted precloacal pores in females (6 vs absent in C. raglai); differs from C. sangi by having a larger size (SVL 73.4–80.9 mm vs 49.9–56.3 mm in C. sangi), more enlarged femoral scales (5–11 vs 4 in C. sangi), fewer precloacal pores in males (6 vs 7 in C. sangi), and more pitted precloacal pores in females (6 vs 4 in C. sangi); differs from C. takouensis by having more enlarged femoral scales (5–11 vs 3–5 in C. takouensis), more precloacal pores in males (6 vs 3–4), the presence of pitted precloacal pores in females (6 vs absent in C. takouensis), and the different dorsal color pattern (irregular bands vs banded in C. takouensis); differs from C. taynguyenensis by having more ventral scale rows (26–38 vs 42–49 in C. taynguyenensis), the presence of enlarged femoral scales (5–11 vs absent in C. taynguyenensis), the different dorsal color pattern (irregular bands vs blotched in C. taynguyenensis), and the presence of transversely enlarged subcaudals (vs absent in C. taynguyenensis); differs from C. yangbayensis by having the presence of pitted precloacal pores in females (6 vs absent in C. yangbayensis), more subdigital lamellae under the fourth toe (17–22 vs 15–17), and fewer subdigital lamellae on first toe (11–14 vs 18–20 in C. yangbayensis); differs from C. ziegleri by having a smaller size (SVL 73.4–80.9 mm vs 84.6–93.0 mm in C. ziegleri), the presence of transversely enlarged subcaudals (vs absent in C. ziegleri), and dark-brown transverse bands of the tail narrower than the light-brown interspaces (vs dark transverse bands wider than the light interspaces in C. ziegleri). Differs from C. chumuensis sp. nov. by having a larger size (SVL 73.4–80.9 mm vs maximum 67.5 mm in C. chumuensis), fewer ventral scale rows (26–38 vs 43–45 in C. chumuensis), and the presence of transversely enlarged subcaudals (vs absent in C. chumuensis). Distribution Cyrtodactylus arndti sp. nov. is currently known only from the Van Canh District, Binh Dinh Province, Vietnam (Fig. 1). Natural history Specimens were found at night between 19:00 and 22:00, on trees or on granite rock, along rocky streams, about 0.6–1.5 m above the ground, at elevations between 150 and 300 m a.s.l. The surrounding habitat was evergreen forest of medium and small hardwoods mixed with shrubs and vines (Fig. 9). The humidity was approximately 40–62% and the air temperature ranged from 28.9 to 33.1 ° C. Other reptiles species found at the sites included Acanthosaura coronata (Günther, 1861) Dixonius vietnamensis Das, 2004, Gekko gecko (Linnaeus, 1758), Gekko sp., Eutropis multifasciata (Kuhl, 1820), Boiga jaspidea (Duméril, Bibron & Duméril, 1854), Psammodynastes pulverulentus (Boie, 1827), and Trimeresurus sp., Published as part of Ngo, Hanh Thi, Hormann, Helene, Pham, Cuong The, Phung, Trung My, Ostrowski, Sabrina, Nguyen, Truong Quang & Ziegler, Thomas, 2023, The discovery of two new species in the Cyrtodactylus irregularis group highlights that hidden diversity remains in the largest clade of the mega-diverse genus Cyrtodactylus, pp. 70-100 in European Journal of Taxonomy 875 (1) on pages 84-94, DOI: 10.5852/ejt.2023.875.2141, http://zenodo.org/record/8064396, {"references":["Ziegler T., Rosler H., Herrmann H. - W. & Vu T. N. 2002. Cyrtodactylus phongnhakebangensis sp. n., ein neuer Bogenfingergecko aus dem annamitischen Karstwaldmassiv, Vietnam. Herpetofauna 24 (141): 11 - 25.","Nguyen S. N., Orlov N. L. & Darevsky I. S. 2006. Descriptions of two new species of the genus Cyrtodactylus Gray, 1827 (Squamata: Sauria: Gekkonidae) from southern Vietnam. Russian Journal of Herpetology 13 (3): 215 - 226. https: // doi. org / 10.30906 / 1026 - 2296 - 2006 - 13 - 3 - 215 - 226","Nguyen S. N., Yang J-X., Le T-N. T., Nguyen L. T., Orlov N., Hoang C. V., Nguyen T. Q., Jin J-Q., Rao D-Q., Hoang T. N., Che J., Murphy R. W. & Zhang Y-P. 2014. DNA barcoding of Vietnamese bent-toed geckos (Squamata: Gekkonidae: Cyrtodactylus) and the description of a new species. Zootaxa 3784 (1): 48 - 66. https: // doi. org / 10.11646 / zootaxa. 3784.1.2","Nguyen S. N., Zhou W-W., Le T-N. T., Tran A-D. T., Jin J-Q., Vo B. D., Nguyen L. T., Nguyen T. T., Nguyen T. Q., Hoang D. D., Orlov N. L., Che J., Murphy R. W. & Zhang Y-P. 2017 a. Cytonuclear discordance, cryptic diversity, complex histories, and conservation needs in Vietnamese bent-toed geckos of the Cyrtodactylus irregularis species complex. Russian Journal of Herpetology 24 (2): 133 - 154.","Nguyen S. N., Nguyen L. T., Nguyen V. D. H. & Murphy R. W. 2017 b. Genetic diversity, phylogeny, and altitudinal distribution of geckos genus Cyrtodactylus on Hon Ba Mountain, Khanh Hoa Province, with note on the subcaudals of C. yangbayensis. Journal of Biotechnology 15 (3 A): 55 - 62.","Heidrich A., Rosler H., Vu N. T., Bohme W. & Ziegler T. 2007. Another new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha-Ke Bang National Park, central Truong Son, Vietnam. Zootaxa 1445: 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1445.1.3","Orlov N. L., Nguyen T. Q., Nazarov A. R., Ananjeva B. N. & Nguyen S. N. 2007. A new species of the genus Cyrtodactylus Gray, 1827 and redescription of Cyrtodactylus paradoxus (Darevsky et Szczerbak, 1997) [Squamata: Sauria: Gekkonidae] from South Vietnam. Russian Journal of Herpetology 14: 145 - 152.","Nazarov R. A., Orlov N. L., Nguyen N. S. & Ho T. C. 2008. Taxonomy of naked-toe geckos Cyrtodactylus irregularis complex of South Vietnam and description of a new species from Chu Yang Sin Natural Park (Krong Bong District, Dac Lac Province), Vietnam. Russian Journal of Herpetology 15: 141 - 156.","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. Zootaxa 3302 (1): 1 - 24. https: // doi. org / 10.11646 / zootaxa. 3302.1.1","Ngo T. V. & Bauer A. M. 2008. Descriptions of two new species of Cyrtodactylus Gray 1827 (Squamata: Gekkonidae) endemic to southern Vietnam. Zootaxa 1715: 27 - 42. https: // doi. org / 10.11646 / zootaxa. 1715.1.2","Ngo T. V., Grismer L. L. & Grismer J. L. 2010. A new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) in Phu Quoc National Park, Kien Giang Biosphere Reserve, Southwestern Vietnam. Zootaxa 2604 (1): 37 - 51. https: // doi. org / 10.11646 / zootaxa. 2604.1.3","Rosler H., Vu T. N., Nguyen T. Q., Ngo V. T. & Ziegler T. 2008. A new Cyrtodactylus (Squamata: Gekkonidae) from central Vietnam. Hamadryad 32: 125 - 140 -","Ngo T. V. & Chan K. O. 2010. A new species of Cyrtodactylus Gray, 1826 (Squamata: Gekkonidae) from Khanh Hoa province, southern Vietnam. Zootaxa 2504 (1): 47 - 60. https: // doi. org / 10.11646 / zootaxa. 2504.1.4","Ngo T. V. & Chan O. K. 2011. A new krastic cave-dwelling Cyrtodactylus Gray (Squamata: Gekkonidae) from Northern Vietnam. Zootaxa 3125 (1): 51 - 63. https: // doi. org / 10.11646 / zootaxa. 1909.1.4","Ngo T. V. & Grismer L. L. 2012. A new species of Cyrtodactylus Gray (Squamata: Gekkonidae) from Tho Chu Island, southwestern Vietnam. Zootaxa 3228 (1): 48 - 60. https: // doi. org / 10.11646 / zootaxa. 3228.1.2","Ziegler T., Nazarov R., Orlov N., Nguyen T. Q., Vu T. N., Dang K. N., Dinh T. H. & Schmitz, A. 2010. A third new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha-Ke Bang National Park, Truong Son Range, Vietnam. Zootaxa 2413 (1): 20 - 36. https: // doi. org / 10.11646 / zootaxa. 2413.1.2","Ziegler T., Phung T. M., Le M. D. & Nguyen T. Q. 2013. A new Cyrtodactylus (Squamata: Gekkonidae) from Phu Yen Province, southern Vietnam. Zootaxa 3686 (4): 432 - 446. https: // doi. org / 10.11646 / zootaxa. 3686.4.2","Luu V. Q., Nguyen T. Q., Do Q. H. & Ziegler T. 2011. A new Cyrtodactylus (Squamata: Gekkonidae) from Huong Son limestone forest, Hanoi, northern Vietnam. Zootaxa 3129: 39 - 50. https: // doi. org / 10.11646 / zootaxa. 3129.1.3","Luu V. Q., Dung T. V., Nguyen T. Q., Le M. D. & Ziegler T. 2017. A new species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from Gia Lai Province, Central Highlands of Vietnam. Zootaxa 4362 (3): 385 - 404. https: // doi. org / 10.11646 / zootaxa. 4362.3.4","Phung T. M., van Schingen M., Ziegler T. & Nguyen T. Q. 2014. A third new Cyrtodactylus (Squamata: Gekkonidae) from Ba Den Mountain, Tay Ninh Province, southern Vietnam. Zootaxa 3764 (3): 347 - 363. https: // doi. org / 10.11646 / zootaxa. 3764.3.5","Le D. T., Nguyen T. Q., Le M. D. & Ziegler T. 2016. A new species of Cyrtodactylus (Squamata: Gekkonidae) from Ninh Binh Province, Vietnam. Zootaxa 4162 (2): 268 - 282. https: // doi. org / 10.11646 / zootaxa. 4162.2.4","Ostrowski S., Le M. D., Ngo H. T., Pham C. T., Nguyen T. Q. & Ziegler T. 2020. A new species of Cyrtodactylus (Squamata: Gekkonidae) from southern Vietnam. Zootaxa 4789 (1): 171 - 203. https: // doi. org / 10.11646 / zootaxa. 4789.1.5","Ostrowski S., Le M. D., Ngo H. T., Pham C. T., Phung T. M., Nguyen T. Q. & Ziegler T. 2021. A new Cyrtodactylus (Squamata: Gekkonidae) from Binh Thuan Province, southern Vietnam. European Journal of Taxonomy 731 (1): 47 - 70. https: // doi. org / 10.5852. ejt. 2021.731.1023","Do Q. H., Phung T. M., Ngo H. T., Le M. D., Ziegler T., Pham C. T. & Nguyen T. 2021. A new species of Cyrtodactylus (Squamata: Gekkonidae) from Ninh Thuan Province, southern Vietnam. Zootaxa 4999 (1): 58 - 76. https: // doi. org / 10.11646 / zootaxa. 4999.1.4"]}

Cyrtodactylus chumuensis sp. nov. urn:lsid:zoobank.org:act: F6A019B5-B1ED-4976-8C1D-803CC0D91DD9 Figs 3–5; Table 2 Diagnosis The new species can be distinguished from remaining congeners of the irregularis species group by a combination of the following characters: maximum SVL 67.5 mm; dorsal pattern with 6 irregularly shaped and short longitudinal stripes on the neck; nuchal band thin, interrupted, reaching the posterior margin of the orbits; the absence of transversely enlarged median subcaudal scales; 4 or 5 enlarged femoral scales on each thigh, 17–19 distinctly enlarged precloacal scales; males with 0 or 1 femoral pore on each thigh, 6 or 7 precloacal pores in a continuous series, ˄-shaped; ventral scales 43–45; dorsal tubercles in 20 irregular longitudinal rows; precloacal groove absent; internasal scales 2; supralabials 8–14; infralabials 9–11; number of subdigital lamellae on fourth finger 16–19 and on fourth toe 19–21. Etymology The new species is named after its type locality, Chu Mu Mountain in Dak Lak Province. We propose the following common names: Chu Mu Bent-toed Gecko (English), Thạch sùng ngón chư mư (Vietnamese). Type material (Figs 3–5) Holotype VIETNAM • ♂; Dak Lak Province, M'Drak District, Ea M’Doal Commune, Chu Mu Mountain; 12°41.330′ N, 108°55.450′ E; 500 m a.s.l.; 20 Jun. 2014; T.M. Phung leg.; Field No. PMT01; IEBR R.4928. Paratypes VIETNAM • 1 ♂ (subadult); Dak Lak Province, M'Drak District, Ea M’Doal Commune, Chu Mu Mountain; 12°41.321′ N, 108°55.382′ E; 400 m a.s.l.; 20 Jun. 2014; T.M. Phung leg.; Field No. PMT02; IEBR R.4929. Description of holotype Adult male; snout-vent length SVL 67.5 mm; tail regenerated 51.4 mm in length (regenerated portion 43.7 mm); body slender, elongate (AG/SVL 0.4); head distinct from neck, elongate (HL/SVL 0.26), relatively wide (HW/HL 0.71) and depressed (HH/HL 0.36); loreal region concave; snout long, blunt in dorsal profile (SE/HL 0.44), longer than diameter of orbit (OD/SE 0.56); scales on snout small, round or oval, granular, larger than scales on occiput; orbit large (OD/HL 0.24), pupils vertical; ear opening small, oval (ED/HL 0.06); rostral wider than high, indented medially the top, in contact with first supralabial, naris and nasorostral on each side, two internasal scales; nostril opening small, oval, surrounded by rostral, nasorostral, two supranasals and one or two postnasals; mental scale triangular, wider than high (ML/MW 0.74); two enlarged, triangular postmentals; supralabials 8/9; infralabials 9/9. Dorsal scales granular, dorsal tubercles round, keeled, conical, in 20 irregular rows at midbody; tubercles on occiput small; tubercles surrounded by 10–12 granular scales; ventral scales smooth, round, midventral scales three times as large as dorsal granular scales, in 45 longitudinal rows at midbody between ventrolateral folds; precloacal groove absent; enlarged femoral scales 4 or 5 on each thigh, the third bearing a femoral pore (Fig. 4C); enlarged precloacal scales 17, arranged in a rhombus; precloacal pores 6, arranged in a ˄-shaped series. Fore and hindlimbs moderately slender (ForeaL/SVL 0.14 mm, CrusL/SVL 0.17); forelimbs dorsally covered by several slightly enlarged tubercles; dorsal hindlimbs with well-developed tubercles; two postcloacal tubercles on each side on the hemipenal swellings; phalanges without webbing; each claw sheathed by two scales, the ventral sheath larger than the upper; number of subdigital lamellae on first finger 11/11, on first toe 10/11, on fourth finger 17/17, on fourth toe 19/19. Coloration in preservative Dorsal surface of head, body and limbs light-brown with some dark-brown bands, pattern without light bordering; occiput marbled with small, irregular dark-brown blotches; rostral, mental and infralabials creamy white, supralabials dark-beige with short greyish brown vertical stripes; neck bands dark-brown, extending in two thin stripes along lateral sides of the snout to the orbits, broader on the neck, interrupted on the left side; two dark-brown longitudinal stripes, disconnected from the neck band extending to shoulders, one dark-brown blotch next to each stripe; dorsal pattern consisting of 6 irregular bands, each formed by two triangularly shaped blotches, shifted medially along the body axis; dorsolateral region covered with small, irregular dark-brown blotches arranged in a longitudinal row from neck to groin; a blurry dark-brown transverse band on dorsal surface of original part of the tail, regenerated part greyish beige and speckled with very small light-greyish brown blotches; dorsal surface of limbs with 3 or 4 dark-brown, blurry bands; phalanges brown with creamy white knuckles; tubercles white or dark-brown depending on position on pattern or background; venter creamy white; ventral tail greyish beige without bands. For coloration of the paratype in life that closely resembles the holotype in life see Fig. 3. Variation The paratype is a subadult and therefore differs greatly in size. Its original tail showed some dark-brown irregular bands, although broken at the base. The number of precloacal pores is 7 and it lacks femoral pores. For more morphological characters see Table 2. Comparisons The new species can be distinguished from all other member of Cyrtodactylus irregularis group from Vietnam by morphological characteristics (see Table 2). Cyrtodactylus chumuensis sp. nov. differs from C. badenensis by having more ventral scale rows (43– 45 vs 25–29 in C. badenensis), the presence of enlarged femoral scales (4–5 vs absent in C. badenensis), the presence of precloacal pores in males (6–7 vs absent in C. badenensis), and the absence of transversely enlarged subcaudals (vs present in C. badenensis); differs from C. bidoupimontis by having a smaller size (SVL 67.5 mm vs 74.0– 86.3 mm in C. bidoupimontis), fewer enlarged femoral scales (4 or 5 vs 8–10 in C. bidoupimontis), a different dorsal color pattern (irregularly banded with longitudinal stripes on the neck vs transversal bands with light borders in C. bidoupimontis), and a thin discontinuous nuchal band (vs well developed, widened posteriorly in C. bidoupimontis); differs from C. bugiamapensis by having fewer enlarged femoral scales (4 or 5 vs 6–10 in C. bugiamapensis) and the different dorsal color pattern (irregularly banded with longitudinal stripes on the neck vs unclear transversal bands formed by irregular round to oblong, dark-brown spots in C. bugiamapensis); differs from C. buchardi by having more ventral scale rows (30 vs 43–45 in C. buchardi), the presence of enlarged femoral scales (4–5 vs absent in C. buchardi), more subdigital lamellae under the fourth finger (16–19 vs 14 in C. buchardi), more subdigital lamellae under the fourth toe (17–21 vs 12 in C. buchardi); differs from C. cattienensis by having more ventral scale rows (43–45 vs 28–42 in C. cattienensis), more subdigital lamellae under the fourth finger (16–19 vs 12–16 in C. cattienensis), and different dorsal color pattern (irregularly banded with longitudinal stripes on the neck vs irregular dark-brown banded, first band on the shoulder x-shaped C. cattienensis); differs from C. caovansungi by having a smaller size (SVL 67.5 mm vs 90.4–94 mm in C. caovansungi), fewer enlarged femoral scales (4 or 5 vs 8 in C. caovansungi), fewer femoral pores on each thigh in males (0–1 vs 6 in C. caovansungi), fewer precloacal pores in males (6 or 7 vs 9 in C. caovansungi), fewer lamellae under the fourth finger (16–19 vs 22 in C. caovansungi), fewer lamellae under the fourth toe (17–21 vs 23–25 in C. caovansungi), and the absence of transversely enlarged subcaudal plates (vs present in C. caovansungi); differs from C. chungi by having more ventral scale rows (43–45 vs 30 or 31 in C. chungi), more dorsal tubercle rows (20 vs 18 in C. chungi), different dorsal color pattern (irregularly banded with longitudinal stripes on the neck vs irregular transversal bands with a closed nuchal band), and a thin, discontinuous nuchal band (vs continuous nuchal band in C. chungi); differs from C. cryptus by having fewer ventral scale rows (43–45 vs 47–50 in C. cryptus), the presence of enlarged femoral scales (vs absent in C. cryptus), fewer precloacal pores in males (6 or 7 vs 9–11 in C. cryptus), a thin, discontinuous nuchal band (vs well developed, widened posteriorly in C. cryptus), and different dorsal color pattern (irregularly banded with short, longitudinal stripes on the neck vs irregular transverse bands in C. cryptus); differs from C. cucdongensis by having more dorsal tubercle rows (20 vs 16–19 in C. cucdongensis), fewer enlarged femoral scales (4 or 5 vs 5–9 in C. cucdongensis), more enlarged precloacal scales (20–21 vs 6–13), and a different dorsal colour pattern (irregularly banded with short, longitudinal stripes on the neck vs irregular dark brown transverse bands); differs from C. culaochamensis by having a smaller size (SVL 67.5 mm vs 69.8–79.8 mm in C. culaochamensis), the presence of enlarged femoral scales (vs absent in C. culaochamensis), fewer lamellae under the first finger (11 vs 13 or 14 in C. culaochamensis), and fewer lamellae under the first toe (10 or 11 vs 13–15 in C. culaochamensis); differs from C. dati by having fewer femoral pores in males (0–2 vs 3 or 4 each side in C. dati), fewer lamellae under the first toe (10 or 11 vs 12 or 13 in C. dati), the presence of blotches on head (vs absent in C. dati), and different dorsal color pattern (irregularly banded with longitudinal stripes on the neck vs irregular dark blotches); differs from C. gialaiensis by the presence of enlarged femoral scales (vs absent in C. gialaiensis), fewer precloacal pores in males (6 or 7 vs 9 or 10 in C. gialaiensis), and more subdigital lamellae under the fourth finger (16–19 vs 14 or 15 in C. gialaiensis) as well as under the fourth toe (17–21 vs 15–17 in C. gialaiensis); differs from C. huynhi by having more dorsal tubercle rows in males (20 vs 16–18 in C. huynhi), fewer lamellae under first finger (11 vs 12–15 in C. huynhi), fewer lamellae under first toe (10 or 11 vs 13–17 in C. huynhi), and a thin discontinuous nuchal band (vs well developed, widened posteriorly in C. huynhi); differs from C. irregularis by having a smaller size (SVL 67.5 mm vs 72.0–86.0 mm in C. irregularis), more ventral scale rows (43–45 vs 37–42 in C. irregularis), fewer enlarged femoral scales (4 or 5 vs 7 or 8 in C. irregularis), and different dorsal color pattern (irregularly banded with short longitudinal stripes on the neck vs blotched in C. irregularis); differs from C. kingsadai by having a smaller size (SVL 67.5 mm vs 83.0–94.0 in C. kingsadai), fewer enlarged femoral scales (4 or 5 vs 9–12 in C. kingsadai), the absence of transversely enlarged subcaudal plates (vs present in C. kingsadai), and more internasals (2 vs 1 in C. kingsadai); differs from C. orlovi by having more ventral scale rows (43–45 vs 36–39 in C. orlovi); a thin, discontinuous nuchal band (vs continuous nuchal band in C. orlovi), and different banded pattern ranges (6 vs 3–5 in C. orlovi); differs from C. phnomchiensis by having a smaller size (SVL 67.5 vs 76.1–80.7 mm in C. phnomchiensis), more precloacal pores in males (6 or 7 vs 5 in C. phnomchiensis), and different dorsal color pattern (irregularly banded vs banded in C. phnomchiensis); differs from C. phuocbinhensis by having a larger size (SVL 67.5 mm vs 46.0– 60.4 mm in C. phuocbinhensis), different dorsal color pattern (irregularly banded vs stripes or blotches in C. phuocbinhensis), and dark-brown transverse banded of the tail than light-brown interspaces (vs dark transverse banded wider than light interspaces in C. phuocbinhensis); differs from C. phumyensis by having more ventral scale rows (43–45 vs 33–41 in C. phumyensis), fewer enlarged femoral scales (4 or 5 vs 5–7 in C. phumyensis), more dorsal tubercle row (20 vs 18 or 19 in C. phumyensis), fewer enlarged precloacal scales (17–19 vs 21–41 in C. phumyensis), and different dorsal color pattern (irregularly banded with short longitudinal stripes on the neck vs anteriorly irregularly spotted and posteriorly banded in C. phumyensis); differs C. pseudoquadrivirgatus by the presence of enlarged femoral scales on each thigh (vs absent in C. pseudoquadrivirgatus), the presence of precloacal pores in males (0–2 vs absent in C. pseudoquadrivirgatus), and more enlarged precloacal scales (17–19 vs 1–12 in C. pseudoquadrivirgatus); differs C. raglai by having a smaller size (SVL 67.5 mm vs 87.5–111.7 mm in C. raglai), more ventral scale rows (43–45 vs 36–39 in C. raglai), fewer enlarged femoral scales (4 or 5 vs 9 or 10 in C. raglai), fewer precloacal pores in males (0–2 vs 5 in C. raglai), and the absence of transversely enlarged subcaudal plates (vs present in C. raglai); differs from C. sangi by having a larger size (SVL 67.5 mm vs 49.9–56.3 mm in C. sangi) and more ventral scale rows (43–45 vs 37 in C. sangi); differs from C. takouensis by having a smaller size (SVL 67.5 mm vs 74.7–81.1 mm in C. takouensis), more ventral scale rows (43–45 vs 39–40 in C. takouensis), more precloacal pores in males (6 or 7 vs 3 or 4 in C. takouensis), the absence of transversely enlarged subcaudal plates (vs present in C. takouensis), a thin discontinuous nuchal band (vs well developed, widened posteriorly in C. takouensis), and different dorsal color pattern (irregularly banded vs banded in C. takouensis); differs from C. taynguyenensis by the presence of enlarged femoral scales on each thigh (vs absent in C. taynguyenensis); irregularly banded of the tail (vs banded in C. taynguyenensis), and different dorsal color pattern (irregularly banded vs blotched in C. taynguyensis); differs from C. yangbayensis by having a smaller size (SVL 67.5 vs 78.5–92.3 mm in C. yangbayensis), more subdigital lamellae under the fourth toe (17–21 vs 15–17 in C. yangbayensis), fewer subdigital lamellae under the first toe (10–11 vs 18–20 in C. yangbayensis), and the absence of transversely enlarged subcaudal plates (vs present in C. yangbayensis); differs from C. ziegleri by having a smaller size (SVL 67.5 vs 84.6–93.0 mm in C. ziegleri), more ventral scale rows (43–45 vs 33–39 in C. ziegleri), and fewer enlarged femoral scales (4 or 5 vs 8–10 C. ziegleri). Distribution Cyrtodactylus chumuensis sp. nov. is currently known only from the Chu Mu Mountain, M'Drak District, Dak Lak Province, Vietnam (Fig. 1). Natural history Specimens were found at night between 19:00 and 22:00, on granite rock, along a rocky stream, approximately 0.5–1.0 m above the ground, at elevations between 400 and 500 m a.s.l. The surrounding habitat was evergreen forest of medium and small hardwoods mixed with shrubs and vines (Fig. 5). The humidity was approximately 50–71% and the air temperature ranged from 27.5 to 32.1°C. Other reptile species found at the sites included Gekko gecko (Linnaeus, 1758), Hemidactylus platyurus (Schneider, 1792), Ahaetulla prasina (Boie, 1827), Lycodon sp., and Oligodon sp., Published as part of Ngo, Hanh Thi, Hormann, Helene, Pham, Cuong The, Phung, Trung My, Ostrowski, Sabrina, Nguyen, Truong Quang & Ziegler, Thomas, 2023, The discovery of two new species in the Cyrtodactylus irregularis group highlights that hidden diversity remains in the largest clade of the mega-diverse genus Cyrtodactylus, pp. 70-100 in European Journal of Taxonomy 875 (1) on pages 78-84, DOI: 10.5852/ejt.2023.875.2141, http://zenodo.org/record/8064396

Ngo, Hanh Thi, Hormann, Helene, Pham, Cuong The, Phung, Trung My, Ostrowski, Sabrina, Nguyen, Truong Quang, Ziegler, Thomas (2023): The discovery of two new species in the Cyrtodactylus irregularis group highlights that hidden diversity remains in the largest clade of the mega-diverse genus Cyrtodactylus. European Journal of Taxonomy 875 (1): 70-100, DOI: https://doi.org/10.5852/ejt.2023.875.2141, URL: http://dx.doi.org/10.5852/ejt.2023.875.2141

Gekko khunkhamensis sp. nov. (Figs. 2���4) Holotype. VNUF R.2021.23 (Field no. SL. KM.23), adult male, (18��11��� N, 104��30��� E, at an elevation of 172 m a.s.l.), Nahin Village, Khunkham District, Khammouane Province, central Laos, collected by S. Sitthivong and Lo O. V. on 29 October 2019. Paratypes. NUOL R.2021.22 (Field no. SL. KM.22), adult female; VNUF R.2021.02 ( Field no. SL. KM.02), adult female, the same data as the holotype. Diagnosis. Gekko khunkhamensis sp. nov. is a member of the Gekko (Japonigekko) japonicus group and differs from all congeners by a combination of the following characters: a medium-sized gecko species (SVL 69.7���75.2 mm); nares in contact with rostral; internasals absent; postmentals enlarged; interorbital scales between anterior corners of the eyes 31 or 32; dorsal tubercles absent; ventrals between mental and cloacal slit 181���185; midbody scales 127���138; ventral scales 42���45; subdigital lamellae on first toe 13 or 14, on fourth toe 14 or 15; tubercles on upper surface of fore and hind limbs absent; precloacal pores absent in the male and females; postcloacal tubercles two; tubercles absent on dorsal surface of tail base; subcaudals distinctly enlarged; dorsal surface of body with five dark grey bands, which become irregular posteriorly. Description of holotype. An adult male with a total length of 158.1 mm (SVL 69.7 mm, TaL 88.4 mm); body slender, elongate (TrunkL/SVL ratio 0.46); head longer than wide (HW/HL ratio 0.75); rostral quadrangular, nearly twice wider than high (RH/RW ratio 0.50) and wider than mental (MW/RW ratio 0.67), in contact with first supralabial and supranasal on each side; nostrils round, in contact with rostral, first supralabial, supranasal, and two enlarged nasals posteriorly; posterior nasal region concave; internasal absent; preorbitals 18/18, preorbital region deeply concave; interorbitals 31; eye large (OD/HL ratio 0.23), pupil vertical; ear opening oval, oblique, smaller than eye (EarL/OD ratio 0.53); mental triangular, wider than long (ML/MW ratio 0.81); enlarged postmentals two, hexagonal, twice longer than wide, and longer than length of mental, touching mental, first infralabial on both sides and seven gular scales posteriorly, outer gular scales larger than inner scales; supralabials 10/10; infralabials 9/9; dorsal scales on body smooth, round or oval, granular, juxtaposed; lateral fold distinct; ventrals distinctly larger than dorsal scales, smooth, imbricate, and largest in the middle of belly; ventrals at midbody 42; scales around midbody 127; ventrals in a row between mental and cloacal slit 185; scales on dorsal surface of forelimbs slightly enlarged; tubercles on dorsal surface of limbs absent; scales on anterior and ventral parts of femur larger than those on posterior and dorsal parts; enlarged femoral scales absent; fingers and toes basally webbed; subdigital lamellae under first fingers 13/13, under fourth fingers 15/15, under first toes 13/13, under fourth toes 15/15; precloacal pores absent; postcloacal tubercles 2/2; base of tail thickened, without tubercles on dorsal surface; subcaudals enlarged, smooth, imbricate. Coloration in life. Dorsal head dark grey with grey blotches; labials grey with light bars; neck with a grey bar; dorsal surface of body grey with five dark grey bands, which become irregular posteriorly; dorsal surface of fore and hind limbs grey with dark blotches; ventral surface of head, belly, and limbs cream with black spots; upper tail with eight grey transverse bands, more distinct posteriorly; underside of tail grey. Sexual dimorphism and variation. Compared to the male holotype, the two female paratypes have a larger size (SVL 71.6���75.2 mm versus 69.7 mm in the male) and more scale rows at midbody (136���138 versus 127 in the male holotype). The dorsal pattern of the paratype (VNUF R.2021.02) is somewhat irregurlarly banded. Measurements and scalation of the female paratypes are shown in Table 2. Comparisons. Based on examination of specimens and data obtained from the literature (Boulenger 1907; Ota et al. 1995; R��sler et al. 2005, 2010, 2011; Yang et al. 2012, Nguyen et al. 2013; Luu et al. 2014, 2015; Ngo et al. 2015; Yang 2015; Luu et al. 2017) we compared the new species from Laos with the remaining members of the Gekko japonicus group (r��sler et al. 2011; Luu et al. 2015, 2017) in Table 3. Below we compared the new species with the closely related species of the G. japonicus group as follows: Gekko khunkhamensis sp. nov. is morphologically most similar to G. tawaensis Okada, 1956. However, the new species can be distinguished from G. tawaensis by having fewer supralabials and infralabials (9 or 10 and 9 or 10 versus 15 and 13, respectively), internasals absent (versus present), postmentals enlarged (versus not enlarged), more subdigital lamellae under first and fourth toes (13 or 14 and 14 or 15 versus 10 and 12, respectively), more postcloacal tubercles (2 versus 1), and marking on upper head present (versus absent). Gekko khunkhamensis sp. nov. does not have dorsal tubercles and thus differs from the following species which have tubercles on the back: G. adleri Nguyen, Wang, Yang, Lehmann, Le, Ziegler & Bonkowski, 2013, G. auriverrucosus Zhou & Liu in Zhou et al., 1982, G. canhi R��sler, Nguyen, Doan, Ho, Nguyen & Ziegler, 2010, G. chinensis Gray, 1842, G. japonicus (Schlegel, 1836), G. hokouensis Pope, 1928, G. kwangsiensis Yang, 2015, G. liboensis Zhou & Li in Zhou et al., 1982, G. palmatus Boulenger, 1907, G. scabridus Liu & Zhou in Zhou et al., 1982, G. shibatai Toda, Sengoku, Hikida & Ota, 2008, G. similignum Smith, 1923, G. swinhonis G��nther, 1864, G. vertebralis Toda, Sengoku, Hikida & Ota, 2008, and G. wenxianensis Zhou & Wang, 2008. Distribution. Gekko khunkhamensis sp. nov. is currently known only from the type locality in Khunkham District, Khammouane Province, central Laos (Fig. 5). Etymology. The specific epithet ��� khunkhamensis ��� refers to the type locality of the new species, Khunkham District of Khammouane Province. We suggest the following common names: Khunkham Gecko (English, German), Kap Ke Khunkham (Laotian) and Gecko de Kunkham (French). Natural history. Specimens of the new species were collected between 19:10 and 20:30, on karst cliffs near the entrance of a cave, from 0.5 to 2.0 m above the ground, at elevations between 172 and 320 m a.s.l. (Fig. 6). The surrounding habitat was disturbed secondary forest of small wood trees and shrubs on the karst cliff. The humidity was approximately 60���70% and the air temperature ranged from 22���26 oC., Published as part of Sitthivong, Saly, Lo, Oanh Van, Nguyen, Truong Quang, Ngo, Hanh Thi, Khotpathoom, Thananh, Le, Minh Duc, Ziegler, Thomas & Luu, Vinh Quang, 2021, A new species of the Gekko japonicus group (Squamata: Gekkonidae) from Khammouane Province, central Laos, pp. 553-571 in Zootaxa 5082 (6) on pages 556-562, DOI: 10.11646/zootaxa.5082.6.3, http://zenodo.org/record/5797638, {"references":["Boulenger, G. A. (1907) Description of new lizards in the British Museum. Annals and Magazine of Natural History, Series 7, 19 (7), 486 - 489. https: // doi. org / 10.1080 / 00222930709487278","Ota, H., Lau, M. W., Weidenhofer, T., Yasukawa, Y. & Bogadek, A. (1995) Taxonomic review of the geckos allied to Gekko chinensis Gray 1842 (Gekkonidae, Reptilia) from China and Vietnam. Tropical Zoology, 8, 181 - 196. https: // doi. org / 10.1080 / 03946975.1995.10539278","Rosler, H., Ziegler, T., Vu, N. T., Herrmann, H. W. & Bohme, W. (2005 \" 2004 \") A new lizard of the genus Gekko Laurenti, 1768 (Squamata: Sauria: Gekkonidae) from the Phong Nha-Ke Bang National Park, Quang Binh Province, Vietnam. Bonner zoologische Beitrage, 53 (1 / 2), 135 - 148.","Rosler, H., Nguyen, Q. T., Doan, V. K., Ho, T. C., Nguyen, T. T. & Ziegler, T. (2010) A new species of the genus Gekko Laurenti (Squamata: Sauria: Gekkonidae) from Vietnam with remarks on G. japonicus (Schlegel). Zootaxa, 2329 (1), 56 - 68. https: // doi. org / 10.11646 / zootaxa. 2329.1.4","Rosler, H., Bauer, A. M., Heinicke, M., Greenbaum, E., Jackman, T., Nguyen, Q. T. & Ziegler, T. (2011) Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae). Zootaxa, 2989 (1), 1 - 50. https: // doi. org / 10.11646 / zootaxa. 2989.1.1","Yang, J. - H., Wang, Y. - Y., Zhang, T. - D., Sun, Y. - J. & Lin, S. - S. (2012) Genetic and morphological evidence on the species validity of Gekko melli Vogt, 1922 with notes on its diagnosis and range extension (Squamata: Gekkonidae). Zootaxa, 3505 (1), 67 - 74. https: // doi. org / 10.11646 / zootaxa. 3505.1.5","Nguyen, Q. T., Wang, Y. - Y. Yang, J. - H., Lehmann, T., Le, M. D., Ziegler, T. & Bonkowski, M. (2013) A new species of the Gekko japonicus group (Squamata: Sauria: Gekkonidae) from the border region between China and Vietnam. Zootaxa, 3652 (5), 501 - 518. https: // doi. org / 10.11646 / zootaxa. 3652.5.1","Luu, V. Q., Calame, T., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2014) A new species of the Gekko japonicus group (Squamata: Gekkonidae) from central Laos. Zootaxa, 3895 (1), 73 - 88. https: // doi. org / 10.11646 / zootaxa. 3895.1.4","Luu, V. Q., Calame, T., Nguyen, T. Q., Le, M. D. & Ziegler, T. (2015) Morphological and molecular review of the Gekko diversity of Laos with descriptions of three new species. Zootaxa, 3986 (3), 279 - 306. https: // doi. org / 10.11646 / zootaxa. 3986.3.2","Ngo, V. T., Pham, H. T., Phimvahan, A., David, P. & Teynie, A. (2015) Gekko aaronbaueri, a new gecko (Squamata: Gekkonidae) from central Laos. Zootaxa, 3914 (2), 144 - 156. https: // doi. org / 10.11646 / zootaxa. 3914.2.4","Yang, J. - H. (2015) A new species of the genus Gekko Laurenti (Squamata: Sauria: Gekkonidae) from Guangxi, China. Zootaxa, 3936 (2), 287 - 295. https: // doi. org / 10.11646 / zootaxa. 3936.2.9.","Luu, V. Q., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2017) A new karst dwelling species of the Gekko japonicus group (Squamata: Gekkonidae) from central Laos. Zootaxa, 4263 (1), 179 - 193. https: // doi. org / 10.11646 / zootaxa. 4263.1.10","Okada, Y. (1956) A new species of gekko from Shikoku, Japan. Annotationes Zoologicae Japonensis, 29, 239 - 241.","Gray, J. E. (1842) Description of some new species of reptiles, chiefly from the British Museum collection. Zoological Miscellany, 2, 57 - 59.","Schlegel, H. (1836) Le Platydactyle Monarque. Platydactylus Monarchus. In: Dumeril, A. M. C. & Bibron, G. (1836) Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Vol. 3. Roret, Paris, pp. 335 - 336.","Pope, C. H. (1928) Four new snakes and a new lizard from South China. American Museum Novitates, 325, 1 - 4.","Toda, M., Sengoku, S., Hikida, T. & Ota, H. (2008) Description of two new species of the genus Gekko (Squamata: Gekkonidae) from the Tokara and Amami Island Groups in the Ryukyu Archipelago, Japan. Copeia, 2008, 452 - 466. https: // doi. org / 10.1643 / CH- 06 - 281","Smith, M. A. (1923) On a collection of reptiles and batrachians from the island of Hainan. Journal of the Natural History Society of Siam, 6, 195 - 212.","Gunther, A. (1864) The Reptiles of British India. The Ray Society, London, xxvii + 452 pp., 26 pls.","Zhou, K. & Wang, Q. (2008) New species of Gekko (Squamata: Sauria: Gekkonidae) from China: morphological and molecular evidence. Zootaxa, 1778 (1), 59 - 68. https: // doi. org / 10.11646 / zootaxa. 1778.1.5"]}

Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang, Ziegler, Thomas (2021): A morphological and molecular review of the genus Goniurosaurus, including an identification key. European Journal of Taxonomy 751 (1): 38-67, DOI: https://doi.org/10.5852/ejt.2021.751.1379, URL: http://dx.doi.org/10.5852/ejt.2021.751.1379

Goniurosaurus lichtenfelderi (Mocquard, 1897) Fig. 5 Diagnosis Body robust; SVL 80.6���113.5 mm; external nares bordered by 5���10 nasal scales; supraorbital region with a row of slightly enlarged tubercles; outer surface of upper eyelid composed of granular scales, about one-half the size of those on top of head and without enlarged tubercles; internasals 1���5 (rarely 1:2; 2:2 or 2:3); supralabials 7���10; infralabials 6���9; preorbital scales 12���19; eyelid fringe scales 43���58; postmentals 2���6; paravertebral tubercles 22���33; scale rows around midbody 117���131, granular scales surrounding tubercles 10���13; axillary pockets shallow; subdigital lamellae under fourth toe 17���21; precloacal pores in males 25���33, in females 17���21; dorsal ground color of head, body and limbs dark purple-brown, without small dark brown blotches; transverse body bands 4, nuchal loop thin, posteriorly rounded, in U-shape; dorsal body bands between limb insertions 2, thin, light yellow; gular region and without dark spots; ventral surfaces of head, body and limbs dull white with a few dark dots on margin regions (modified after Grismer 2000; Grismer et al. 2002; Nguyen 2011). Description (Supp. file 1: Table S2) Body robust, adult males: SVL 80.6���113.5 mm (mean �� SE: 97.8 �� 0.7 mm, n =92), TaL 6.2��� 84.8 mm (61.5�� 1.7 mm); adult females: SVL 81.0��� 105.5 mm (96.0�� 0.7 mm, n=72), TaL 22.7��� 81.2 mm (58.4 �� 1.4 mm); juveniles: SVL 41.1���77.1 mm (65.4�� 2.9 mm, n=14), TaL 20.9���64.7 mm (48.3 �� 3.4 mm) (Supp. file 1: Table S2); head triangular, wider than neck, covered by uniform granular scales interspersed with tubercles in temporal and occipital regions; scales on rostrum slightly larger and flatter; enlarged supraorbital tubercles in a conspicuous row; middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial on each side, dorsolaterally by prenasal on each side, and dorsally by 1 or 2 internasals and two supranasals; internasals 1���5 (rarely 1:2; 2:2 or 2:3); external nares bordered by 5���10 nasals; preorbital scales 12���18; supralabials 7���10, grading into granular scales posteriorly; infralabials 6���9; eyes large, pupils vertical; eyelid fringe scales 47���58, those of upper eyelid slightly enlarged; outer surface of upper eyelid composed of granular scales of about one-half the size of those on top of head, without enlarged tubercles; fold of skin originating in the suborbital region extends posteroventrally across angle of jaw; external auditory meatus elliptical; tympanum deeply recessed; mental triangular, bordered laterally by first infralabial on each side and posteriorly by 2���5 postmentals; postmentals bordered by 7���10 gular scales; gular region below lower jaws without enlarged tubercles; gular scales juxtaposed and granular, abruptly grading posteriorly into flat hexagonal scales and even larger ventral scales. Neck narrower than body, covered with uniform granular scales interspersed with several sharply conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat more flat; dorsal body tubercles surrounded by 10���13 granular scales; dorsal tubercle rows at midbody 15���22; paravertebral tubercles between limb insertions 22���33, distinct vertebral row of tubercles absent; scales around midbody 117���130; ventral scales large; males with 24���33 precloacal pores in a transverse continuous series, females without distinct precloacal pores (but 25���33 pitted scales present); region posterior to vent covered by flat juxtaposed scales and greatly swollen, containing 1 (rarely 2) enlarged tubercles on each side at the level of vent; tail thick at base; light band on tail base in width of 4���7 scales and with 6���10 tubercles in a transversal series. Limbs robust, covered dorsally with granular scales interspersed with several tubercles and ventrally with flat juxtaposed to subimbricate scales; dorsal granular scales grade into slightly flattened subimbricate scales on top of pes and manus; hind limbs larger than forelimbs; axillary pockets shallow; subdigital lamellae wide, 6���9 under first finger, 13���17 under fourth finger, 6���10 under first toe, 16���20 under fourth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth (modified after Grismer 2000; Grismer et al. 2002; Nguyen 2011). Coloration in life (Fig. 5) Dorsal ground color of head, body and limbs grey and grey brown in juveniles without blotches and chestnut brown in adults with dark brown blotches (in animals with SVL> 80 mm); iris orange or redish brown; four bands on the dorsal ground, thin, yellow in both juveniles and adults, immaculate without dark spots (few dark brown spots present in some animals), all edged anteriorly and posteriorly by thin dark brown bands, including one thin nuchal loop extending from posterior corners of eyes and posteriorly rounded (in U-shape), two body bands between limb insertions, and another one on tail base; ground color of tail dark brown, and grey brown at mottled tail base; 3���5 immaculate white caudal bands except first band slightly yellow, and some animals with regenerated tail present immaculate grey brown without white caudal bands; ventral surfaces of head, body and limbs dull white with a few dark dots on margin regions of belly, limbs, gular region, but immaculate dull white in juveniles (SVL ��� 65 mm) without dark spots. Morphological comparisons Our morphological comparisons of the tiger gecko species (except for G. araneus) from Vietnam revealed an overall significant difference in the snout-vent length (SVL). As the result, G. huuliensis had the largest SVL, while G. lichtenfelderi had the shortest SVL in both adult males and females, compared to the other tiger geckos (P G. catbaensis was not significantly different from that of G. luii (P> 0.05, Supp. file 2: Fig. S1). The PCA of 18 selected morphometric characters estimated the first (PC1) and second principal (PC2) components to explain 82.5% (74.5% and 8.0%, respectively) of the variance among the four investigated species of Goniurosaurus in Vietnam (Supp. file 2: Fig. S2). The extracted PC1 and PC2 scores of the PCA differ significantly among the four species (Kruskal-Wallis���s test, P G. lichtenfelderi are not concordant with the three remainders (Fig. S2). Three morphological factors, namely head length (HL), head width (HW), and mouth width (MW) highly account for the overall difference in the PC1 score, while the PC2 score is mainly explained by the body height (BH) and body width (BW) (Supp. file 2: Fig. S2). Regarding the comparisons of the 17 species of Goniurosaurus from China and Vietnam, the multiple correspondence analysis (MCA) clustered them separately into three groups (G. lichtenfelderi group, G. luii group and G. yingdeensis group) represented with different meristic spaces on the spatial coordinate of Dim 1 and Dim 2 (Fig. 6A, Table 1). The characters of fourth toe (LT4), body limb shape and body bands highly account for the Dim1 score, whereas the characters of precloacal pores (PP) and granular scales surrounding dorsal tubercles (GST) are the most correlated with the Dim 2 score (Fig. 6B)., Published as part of Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, pp. 38-67 in European Journal of Taxonomy 751 (1) on pages 52-54, DOI: 10.5852/ejt.2021.751.1379, http://zenodo.org/record/4890455, {"references":["Grismer L. L. 2000. Goniurosaurus murphyi Orlov and Darevsky: a junior synonym of Goniurosaurus lichtenfelderi Mocquard. Journal of Herpetology 34 (3): 486 - 488. https: // doi. org / 10.2307 / 1565379","Grismer L. L., Shi H. T., Orlov N. L. & Ananjeva N. B. 2002. A new species of Goniurosaurus (Squamata: Eublepharidae) from Hainan Island, China. Journal of Herpetology 36: 217 - 224. https: // doi. org / 10.2307 / 1565994","Nguyen T. Q. 2011. Systematics, Ecology, and Conservation of the Lizard Fauna in Northeastern Vietnam, with Special Focus on Pseudocalotes (Agamidae), Goniurosaurus (Eublepharidae), Sphenomorphus and Tropidophorus (Scincidae) from this Country. PhD thesis, University of Bonn, Germany."]}

Key to the species of Goniurosaurus (Fig. 7, Table 1) Modified from Grismer et al. (1999, 2002), Vu et al. (2006), Orlov et al. (2008), Ziegler et al. (2008), Wang et al. (2010, 2013, 2014), Nguyen (2011), Chen et al. (2014), Yang & Chan (2015), Honda & Ota (2017), Zhou et al. (2018, 2020a, 2020b), Qi et al. (2020a, 2020b) and the present study. 1. Precloacal pores in males present, claws are sheathed by scales...................................................... 7 ��� Precloacal pores absent, unsheathed claws (G. kuroiwae group)..................................................... 2 2. Yellow brown to gold iris; a single scale at the base of each digit which is occasionally slightly enlarged............................................................................................... G. yamashinae (Okada, 1936) ��� Blood-red iris; one to three enlarged scales at the base of each digit on the limbs.......................... 3 3. Adults without interspace mottling................................................................................................... 4 ��� Adults with interspace mottling........................................................................................................ 5 4. Robust body stature; dorsal body tubercles between the limb insertions in cross-section are triangular to elliptical and sharply keeled anteriorly; 34���42 paravertebral tubercles; ventral scales are juxtaposed and sharply raised..................................... G. toyamai Grismer, Ota & Tanaka, 1994 ��� Slender body stature; dorsal body tubercles between the limb insertions in cross-section are smooth or very weakly keeled; 22���29 paravertebral tubercles; ventrals are flat, wide, and subimbricate to imbricate............................................................................ G. splendens (Nakamura & U��no, 1959) 5. Dorsal banding absent or incomplete................................................... G. kuroiwae (Namiye, 1912) ��� Prominent dorsal pattern consisting of transverse bands between the nape of the neck and the caudal constriction....................................................................................................................................... 6 6. No such reddish or pinkish tint in dorsal pattern and iris......................... G. orientalis (Maki, 1931) ��� Reddish or pinkish tint to some extent in dorsal pattern and iris.............................................................................................................................................................. G. sengokui (Honda & Ota, 2017) 7. Precloacal pores in males less than 16 (G. yingdeensis group)........................................................ 8 ��� Precloacal pores in males more than 16..........................................................................................11 8. Scales around midbody 121���128; longitudinal dorsal tubercle rows at midbody 16���17...................................................................................... G. gollum Qi, Wang, Grismer, Chen, Lyu & Wang, 2020 ��� Scales around midbody 99���115; longitudinal dorsal tubercle rows at midbody 20���28................... 9 9. Nuchal loop and body bands with small dark blotches; enlarged row of supraorbital tubercles absent; trunk of body usually with a longitudinal light vertebral stripe................................................................................................................................. G. varius Qi, Grismer, Lyu, Zhang, Li & Wang, 2020 ��� Nuchal loop and body bands without dark blotches; enlarged row of supraorbital tubercles present; trunk of body without a longitudinal light vertebral stripe............................................................. 10 10. Tubercles between orbits present; gular scales bordering the postmentals 2���4; preorbital scales 5���7............................................................................................ G. yingdeensis Wang, Yang & Cui, 2010 ��� Tubercles between orbits absent; gular scales bordering the postmentals 4���6; preorbital scales 7���9........................................................................................ G. zhelongi Wang, Jin, Li & Grismer, 2014 11. Body and limbs robust (G. lichtenfelderi group)............................................................................ 12 ��� Body and limbs splayed gracile (G. luii group).............................................................................. 16 12. Precloacal pores in males 37���46....... G. bawanglingensis Grismer, Haitao, Orlov & Anajeva, 2002 ��� Precloacal pores in males fewer than 37......................................................................................... 13 13. Nuchal loop protracted posteriorly (in V-shape); number of body bands 4; axillary pockets deep; precloacal pores in males more than 33......................... G. zhoui Zhou, Wang, Chen & Liang, 2018 ��� Nuchal loop rounded posteriorly (in U-shape); number of body bands 3; axillary pockets shallow; precloacal pores in males fewer than 33......................................................................................... 14 14. Eyelid fringe scales 55���70; enlarged row of supraorbital tubercles absent; scales of upper eyelid to top of head equal of those on the top of the head in size.................... G. hainanensis Barbour, 1908 ��� Eyelid fringe scales 47���58; enlarged row of supraorbital tubercles present; scales of upper eyelid to top of head �� of those on the top of the head in size..................................................................... 15 15. Lateral spotting on belly absent; preorbital scales 18���21; scales around midbody 109���118; subdigital lamellae under the fourth finger 18���19.............................................. G. kwanghua Zhu & He, 2020 ��� Lateral spotting on belly present; preorbital scales 12���18; scales around midbody 117���130; subdigital lamellae under the fourth finger 13���17...................................... G. lichtenfelderi (Mocquard, 1897) 16. Internasal absent........................... G. catbaensis Ziegler, Nguyen, Schmitz, Stenke & R��sler, 2008 ��� Internasal present............................................................................................................................ 17 17. Postcloacal tubercles 3���6................................................. G. araneus Grismer, Viets & Boyle, 1999 ��� Postcloacal tubercles 1���3................................................................................................................ 18 18. Adult body length (SVL)> 126 mm; dorsum without dark blotches............................................................................................................... G. huuliensis Orlov, Ryabov, Nguyen, Nguyen & Ho, 2008 ��� Maximum body length (SVL) ��� 126 mm; dorsum with dark blotches.......................................... 19 19. Dorsum with many small dark blotches.................................. G. luii Grismer, Viets & Boyle, 1999 ��� Dorsum with rarely scattered dark blotches................................................................................... 20 20. Outer surface of upper eyelid composed of granular scales, about the same size of those on top of head and with enlarged tubercles.......................................................... G. gezhi Zhu, He & Li, 2020 ��� Enlarged row of supraorbital tubercles present; scales of upper eyelid to top of head �� of those on the top of the head in size............................................................................................................... 21 21. Precloacal pores in males 31���33.............................................. G. kwangsiensis Yang & Chan, 2015 ��� Precloacal pores in males fewer than 31......................................................................................... 22 22. Lateral spotting on belly present; nasal scales 6���7; paravertebral tubercles between limb insertions 30���34; precloacal pores 26���28............................................... G. kadoorieorum Yang & Chan, 2015 ��� Lateral spotting on belly absent, nasal scales 8���9; paravertebral tubercles between limb insertions 27���28; precloacal pores 23.............................................. G. liboensis Wang, Yang & Grismer, 2013, Published as part of Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, pp. 38-67 in European Journal of Taxonomy 751 (1) on pages 58-60, DOI: 10.5852/ejt.2021.751.1379, http://zenodo.org/record/4890455, {"references":["Grismer L. L., Viets B. E. & Boyle L. J. 1999. Two new continental species of Goniurosaurus (Squamata: Eublepharidae) with a phylogeny and evolutionary classification of the genus. Journal of Herpetology 33 (3): 382 - 393. https: // doi. org / 10.2307 / 1565635","Grismer L. L., Shi H. T., Orlov N. L. & Ananjeva N. B. 2002. A new species of Goniurosaurus (Squamata: Eublepharidae) from Hainan Island, China. Journal of Herpetology 36: 217 - 224. https: // doi. org / 10.2307 / 1565994","Vu N. T., Nguyen Q. T., Grismer L. L. & Ziegler T. 2006. First record of the Chinese leopard gecko, Goniurosaurus luii (Reptilia: Eublepharidae) from Vietnam. Current Herpetology 25: 93 - 95. https: // doi. org / 10.3105 / 1345 - 5834 (2006) 25 [93: FROTCL] 2.0. CO; 2","Orlov N. L., Ryabow S. A., Nguyen T. T., Nguyen Q. T. & Ho T. C. 2008. A new species of Goniurosaurus (Sauria: Gekkota: Eublepharidae) from north Vietnam. Russian Journal of Herpetology 15: 229 - 244.","Ziegler T., Nguyen T. Q., Schmitz A., Stenke R. & Rosler H. 2008. A new species of Goniurosaurus from Cat Ba Island, Hai Phong, northern Vietnam (Squamata: Eublepharidae). Zootaxa 1771: 16 - 30. https: // doi. org / 10.11646 / zootaxa. 1771.1.2","Wang Y. Y., Yang J. H. & Cui R. F. 2010. A new species of Goniurosaurus (Squamata: Eublepharidae) from Yingde, Guangdong Province, China. Herpetologica 66: 229 - 240. https: // doi. org / 10.1655 / 09 - 046 r 2.1","Wang Y. Y., Yang J. H. & Grismer L. L. 2013. A new species of Goniurosaurus (Squamata: Eublepharidae) from Libo, Guizhou Province, China. Herpetologica 69: 214 - 226. https: // doi. org / 10.1655 / herpetologica-d- 12 - 00084","Wang Y. Y., Jin M. J., Li Y. L. & Grismer L. L. 2014. Description of a new species of Goniurosaurus (Squamata: Eublepharidae) from the Guangdong Province, China, based on molecular and morphological data. Herpetologica 70: 309 - 322. https: // doi. org / 10.1655 / herpetologica-d- 13 - 00080","Nguyen T. Q. 2011. Systematics, Ecology, and Conservation of the Lizard Fauna in Northeastern Vietnam, with Special Focus on Pseudocalotes (Agamidae), Goniurosaurus (Eublepharidae), Sphenomorphus and Tropidophorus (Scincidae) from this Country. PhD thesis, University of Bonn, Germany.","Chen T. B., Meng Y. J., Jiang K., Li P. P., Wen B. H., Lu W., Lazell J. & Hou M. 2014. New record of the leopard gecko Goniurosaurus araneus (Squamata: Eublepharidae) for China and habitat portioning between geographically and phylogenetically close leopard geckos. IRCF Reptiles & Amphibians 21 (1): 16 - 27.","Yang J. H. & Chan B. P. 2015. Two new species of the genus Goniurosaurus (Squamata: Sauria: Eublepharidae) from southern China. Zootaxa 3980 (1): 67 - 80. https: // doi. org / 10.11646 / zootaxa. 3980.1.4","Honda M. & Ota H. 2017. On the live coloration and partial mitochondrial DNA sequences in the topotypic population of Goniurosaurus kuroiwae orientalis (Squamata: Eublepharidae), with description of a new subspecies from Tokashikijima Island, Ryukyu Archipelago, Japan. Asian Herpetological Research 8: 96 - 107. https: // doi. org / 10.16373 / j. cnki. ahr. 170003","Zhou R. B., Wang N., Chen B. & Liang B. 2018. Morphological evidence uncovers a new species of Goniurosaurus (Squamata: Eublepharidae) from the Hainan Island, China. Zootaxa 4369 (2): 281 - 291. https: // doi. org / 10.11646 / zootaxa. 4369.2.8","Qi S., Grismer L. L., Lyu Z. T., Zhang L., Li P. P. & Wang Y. Y. 2020 a. A definition of the Goniurosaurus yingdeensis group (Squamata, Eublepharidae) with the description of a new species. ZooKeys 986: 127 - 155. https: // doi. org / 10.3897 / zookeys. 986.47989","Qi S., Wang J., Grismer L. L., Chen H-H., Lyu Z-T. & Wang Y-Y. 2020 b. The stoor hobbit of Guangdong: Goniurosaurus gollum sp. nov., a cave-dwelling leopard gecko (Squamata, Eublepharidae) from South China. ZooKeys 991: 137 - 153. https: // doi. org / 10.3897 / zookeys. 991.54935","Zhu X. Y., Chen G. Y., Roman-Palacios C., Li Z. & He Z. Q. 2020 a. Goniurosaurus gezhi sp. nov., a new gecko species from Guangxi, China (Squamata: Eublepharidae). Zootaxa 4852 (2): 211 - 222. https: // doi. org / 10.11646 / zootaxa. 4852.2.6"]}

Goniurosaurus luii Grismer, Viets & Boyle, 1999 Fig. 4 Diagnosis Body splayed and gracile, SVL 86.5���126.5 mm; external nares bordered by 5���8 nasal scales; supraorbital region with a row of enlarged tubercles; outer surface of upper eyelid composed of granular scales, about one half the size of those on top of head and without enlarged tubercles; internasals 1 or 2; supralabials 8���12; infralabials 8���11; preorbital scales 13���16; eyelid fringe scales 46���61; postmentals 2���6; gular region below lower jaws with enlarged tubercles; paravertebral tubercles 29���38; scale rows around midbody 119��� 144, granular scales surrounding tubercles 9���14; axillary pockets deep; subdigital lamellae under fourth toe 20���26; precloacal pores in males 23���32; iris brown or bright orange; dorsal ground color of head, body and limbs pale brown to grey brown, mottled with small dark brown blotched; nuchal loop thin, posteriorly protracted (in V-shape); dorsal body bands between limb insertions 3, thin, immaculate yellow; gular region, belly, and ventral surface of limbs with brown spots (Grismer et al. 1999; Vu et al. 2006; Nguyen 2011). Description (Supp. file 1: Table S2) Body splayed and gracile, males: SVL 88.8���123.0 mm (mean ��SE: 109.3 �� 1.4 mm, n =34), TaL 2.8���96.8 mm (72.5 �� 3.8 mm); adult females: SVL 86.5���126.5 mm (112.4 �� 1.3 mm, n=43), TaL 43.3���102.2 mm (72.2�� 2.1 mm); juveniles: SVL 55.0��� 84.8 mm (74.8 �� 3.1 mm, n=11), TaL 44.2��� 68.9 (58.7 �� 2.7 mm) (Supp. file 1: Table S2); head triangular, wider than neck, covered by uniform granular scales interspersed with tubercles in temporal and occipital regions; scales on rostrum slightly larger and flatter; enlarged supraorbital tubercles in a conspicuous row; middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial on each side, dorsolaterally by prenasal on each side, and dorsally by 1 or 2 internasal and two supranasals; internasals 1 or 2; external nares bordered by 5���8 nasal scales (prenasal, anterior and posterior supranasals, 2 slightly enlarged postnasal, and 1���3 granular scales); preorbital scales 13���16; supralabials 8���12, grading into granular scales posteriorly; infralabials 8���11; eyes large, pupils vertical; eyelid fringe scales 46���56, those of upper eyelid slightly enlarged; outer surface of upper eyelid composed of granular scales, about one half the size of those on top of head and without enlarged tubercles; fold of skin originating in suborbital region extends posteroventrally across angle of jaw; external auditory meatus elliptical with long axis directed dorsoventrally; tympanum deeply recessed; mental triangular, bordered laterally by first infralabial on each side and posteriorly by 2���6 postmentals; postmentals bordered by 6���11 gular scales; gular region below lower jaws with enlarged tubercles; gular scales juxtaposed and granular, abruptly grading posteriorly into flat hexagonal pectoral scales and even larger ventral scales. Neck narrower than body, covered with uniform granular scales interspersed with several sharply pointed conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat more flat; dorsal body tubercles surrounded by 11���13 granular scales; dorsal tubercle rows at midbody 20���24; paravertebral tubercles between limb insertions 29���38, distinct vertebral row of tubercles absent; scale rows around midbody 119���144; ventral scales large; males with 24���32 precloacal pores in a transverse continuous series, females without precloacal pores (but pitted scales present); region posterior to vent covered by flat juxtaposed scales and greatly swollen, containing 1���3 enlarged tubercles on each side at level of vent; tail thick at base, light band on tail base in width of 9���13 granular scales with 9���10 tubercles in transversal series; ventral caudals of tail base larger than dorsal caudals. Limbs relatively long and thin, covered dorsally with granular scales interspersed with several tubercles and ventrally with flat juxtaposed to subimbricate scales; dorsal granular scales grade into slightly flattened subimbricate scales on top of pes and manus; hind limbs larger than forelimbs; axillary pockets deep; subdigital lamellae wide, 9���12 under first finger, 17���22 under fourth finger, 10���12 under first toe, 20���26 under fourth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth (Grismer et al. 1999; Vu et al. 2006; Nguyen 2011). Coloration in life (Fig. 4) Dorsal ground color of head, body and limbs pale brown to grey brown, mottled with small dark brown circle blotches, juveniles without small dark brown blotches (except juveniles SVL ��� 72 mm initially present on head); iris red brown or bright orange; five bands on dorsal ground, thin, immaculate without dark spots, slight bisque in juveniles, orange brown or yellow in adults, all edged anteriorly and posteriorly by thin dark brown bands, including one thin nuchal loop extending from posterior corners of eyes and posteriorly protracted (in V-shape), three body bands between limb insertions, and another one on tail base; ground color of tail nearly solid black, and grey brown at mottled tail base; 3���6 immaculate white caudal bands; ventral surfaces of head, body and limbs dull white with a few dark brown spots in gular region, on belly and limbs., Published as part of Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, pp. 38-67 in European Journal of Taxonomy 751 (1) on pages 50-52, DOI: 10.5852/ejt.2021.751.1379, http://zenodo.org/record/4890455, {"references":["Grismer L. L., Viets B. E. & Boyle L. J. 1999. Two new continental species of Goniurosaurus (Squamata: Eublepharidae) with a phylogeny and evolutionary classification of the genus. Journal of Herpetology 33 (3): 382 - 393. https: // doi. org / 10.2307 / 1565635","Vu N. T., Nguyen Q. T., Grismer L. L. & Ziegler T. 2006. First record of the Chinese leopard gecko, Goniurosaurus luii (Reptilia: Eublepharidae) from Vietnam. Current Herpetology 25: 93 - 95. https: // doi. org / 10.3105 / 1345 - 5834 (2006) 25 [93: FROTCL] 2.0. CO; 2","Nguyen T. Q. 2011. Systematics, Ecology, and Conservation of the Lizard Fauna in Northeastern Vietnam, with Special Focus on Pseudocalotes (Agamidae), Goniurosaurus (Eublepharidae), Sphenomorphus and Tropidophorus (Scincidae) from this Country. PhD thesis, University of Bonn, Germany."]}

Goniurosaurus catbaensis Ziegler, Nguyen, Schmitz, Stenke & R��sler, 2008 Fig. 2 Diagnosis Body splayed and gracile, SVL 89.3���125.3 mm; external nares bordered by 6���8 nasal scales; supraorbital region with a row of enlarged tubercles; outer surface of upper eyelid composed of granular scales, about the same size of those on top of head and with a row of 6���10 enlarged tubercles; internasals absent; supralabials 8���11; infralabials 7���10; eyelid fringe scales 45���56; postmentals 2���5; gular region below lower jaws without enlarged tubercles; paravertebral tubercles 31���38; scale rows around midbody 112��� 127, granular scales surrounding tubercles 8���11; axillary pockets deep; subdigital lamellae under fourth toe 22���25; precloacal pores in males 16���23; iris orange-brown; dorsal ground color of head, body and limbs gray-brown to pale brown and mottled with dark brown blotches; nuchal loop thin, posteriorly protracted (in V-shape); dorsal body bands between limb insertions 3���4, thin, yellow, without dark spots; light band on tail base in width of 8���9 scales; ground color of tail black, caudal bands 5, white, completed ventrally; ventral surface of head, body and limbs dull white and immaculate, gular region with brown spots (modified after Ziegler et al. 2008; Nguyen 2011). Description (Supp. file 1: Table S2) Body splayed and gracile, adult males: SVL 92.9���125.3 mm (mean �� SE: 112.3�� 0.8 mm, n =80), TaL 7.2��� 97.9 mm (75.2 �� 1.9 mm); adult females: SVL 89.3���122.1 mm (111.8�� 0.8 mm, n =93), TaL 17.5��� 98.3 mm (70.3 �� 1.7 mm); juveniles: SVL 53.4���78.8 mm (68.8 �� 1.8 mm, n =21), TaL 11.8���61.2 mm (48.1 �� 2.6 mm) (Supp. file 1: Table S2); head triangular, wider than neck, covered by uniform granular scales interspersed with tubercles in temporal and occipital regions; scales on rostrum slightly larger and flatter; enlarged supraorbital tubercles in a conspicuous row; middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial on each side, dorsolaterally by prenasal on each side, and dorsally by two supranasals; internasal (postrostral) scales absent; external nares bordered by 6���8 nasals: anteriorly by prenasal and supranasal, dorsally by supranasal, posteriorly by two slightly enlarged postnasals and 1 or 2 smaller granular scales, and ventrally by prenasal; prenasals with long recurved ventral portion; supranasals triangular, meeting in midline behind rostral suture; preorbital scales 10���13; supralabials 8���11; infralabials 7���10; eyes relatively large, pupils vertical; eyelid fringe scales 45���56, those of upper eyelid slightly enlarged; outer surface of upper eyelid composed of granular scales of about the same size of those on top of head, including a row of 6���10 enlarged tubercles; fold of skin originating from suborbital region extends posteroventrally across angle of jaw; external auditory meatus elliptical with long axis directed dorsoventrally; tympanum deeply recessed; mental triangular, bordered laterally by first infralabial on each side and posteriorly by 2���5 postmentals; postmentals bordered by 6���10 gular scales; gular region below lower jaws without enlarged tubercles; gular scales juxtaposed and granular, abruptly grading posteriorly into flat hexagonal pectoral scales and even larger ventral scales. Neck narrower than body, covered with uniform granular scales interspersed with several sharply pointed conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat lower in profile; dorsal body tubercles surrounded by 9���11 granular scales; dorsal tubercle rows at midbody 19���24; paravertebral tubercles between limb insertions 31���38, distinct vertebral row of tubercles absent; scales around midbody 112���127; larger ventral scales grade abruptly into smaller granular scales immediately anterior to vent at level of preanal pores; males with 18���23 precloacal pores in a transverse continuous series; region posterior to vent covered by flat juxtaposed scales and great hemipenial bulges, containing 2 or 3 enlarged postcloacal tubercles laterally on each side at level of vent; tail long and thin, thick at base, anteriorly with whorls; dorsal tail scales flat, smooth, up to 1.5 times the size of dorsal scales, arranged in more or less regular transverse rows; light band on tail base in width of 8���9 granular scales and with 7���8 tubercles in a transversal series; subcaudals larger than those on dorsal surface of tail. Limbs relatively long and thin, covered dorsally with granular scales interspersed with several tubercles and ventrally with flat juxtaposed to subimbricate scales; dorsal granular scales grade into slightly flattened subimbricate scales on top of pes and manus; hind limbs slightly larger than forelimbs; larger granular scales on ventral surface of pes and manus; axillary pockets deep; subdigital lamellae wide, 9���11 under first finger, 18���21 under fourth finger, 9���12 under first toe, 22���25 under fourth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth (modified after Ziegler et al. 2008; Nguyen 2011). Coloration in life (Fig. 2) Dorsal ground color of head, body and limbs grey brown (in animals with SVL 53.4���105 mm) and blotches chestnut brown (in animals with SVL 110���125.3 mm), juveniles without small blotches, adults mottled with few circular blotches on body and limbs and long dark brown blotches on head; few yellow conical tubercles on franks of neck and body, and limbs in adults; iris light orange or red brown; five bands on the dorsal ground, thin, immaculate without dark spots, cream in juveniles and light orange or yellow in adults, all edged anteriorly and posteriorly by thin dark brown bands, including one thin nuchal loop extending from posterior corners of eyes and posteriorly protracted (in V-shape), three body bands between limb insertions, and another one on tail base; ground color of tail dark brown, and grey brown at mottled tail base; 3���5 immaculate white caudal bands, edged anteriorly and posteriorly in black; ventral surface of head, body and limbs dull white, juveniles absolutely immaculate, but adults with few dark spots on limbs, weak brown lateral spotting in gular region, venter and limbs., Published as part of Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, pp. 38-67 in European Journal of Taxonomy 751 (1) on pages 45-47, DOI: 10.5852/ejt.2021.751.1379, http://zenodo.org/record/4890455, {"references":["Ziegler T., Nguyen T. Q., Schmitz A., Stenke R. & Rosler H. 2008. A new species of Goniurosaurus from Cat Ba Island, Hai Phong, northern Vietnam (Squamata: Eublepharidae). Zootaxa 1771: 16 - 30. https: // doi. org / 10.11646 / zootaxa. 1771.1.2","Nguyen T. Q. 2011. Systematics, Ecology, and Conservation of the Lizard Fauna in Northeastern Vietnam, with Special Focus on Pseudocalotes (Agamidae), Goniurosaurus (Eublepharidae), Sphenomorphus and Tropidophorus (Scincidae) from this Country. PhD thesis, University of Bonn, Germany."]}

Goniurosaurus huuliensis Orlov, Ryabov, Nguyen, Nguyen & Ho, 2008 Fig. 3 Diagnosis Body splayed and gracile, SVL 97.2���134.6 mm; external nares bordered by 6���8 nasal scales; supraorbital region with a row of enlarged tubercles; outer surface of upper eyelid composed of granular scales, about one half the size of those on top of head and without enlarged tubercles; internasal 1 or 2 (rarely absent); supralabials 9���12; infralabials 9���12; preorbital scales 14���20; eyelid fringe scales 51���59; postmentals 2���4; gular region below lower jaws with enlarged tubercles; paravertebral tubercles 31���37; scale rows around midbody 118���130, granular scales surrounding tubercles 11���13; axillary pockets deep; subdigital lamellae under fourth toe 21���25; precloacal pores in males 25���30; iris reddish brown; dorsal ground color of head, body and limbs dark brown, without small dark brown blotches (dark blotches present only on lower zone of flanks); nuchal loop thin, posteriorly protracted (in V-shape); dorsal body bands between limb insertions 3, thin, immaculate yellow; gular region with brown spots (modified after Orlov et al. 2008; Nguyen 2011). Description (Supp. file 1: Table S2) Body splayed and gracile, adult males SVL 97.2���132.2 mm (mean �� SE: 118.9�� 1.4 mm, n =32), TaL 36.7���108.6 mm (72.7 �� 3.7 mm), adult females SVL 97.4���134.6 mm (121.1 �� 1.2 mm, n =46), TaL 48.4���106.7 mm (78.2 �� 2.7 mm), juveniles SVL 73.5���74.4 mm (n =2), TaL 62.5���62.7 mm (Supp. file 1: Table S2); head triangular, wider than neck, covered by uniform granular scales interspersed with tubercles in temporal and occipital regions; scales on rostrum slightly larger and flatter; enlarged supraorbital tubercles in a conspicuous row; middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial on each side, dorsolaterally by prenasal on each side, and dorsally by internasal and two supranasals; internasal 1���3 (rarely absent); external nares bordered by 5���7 nasals (prenasal, anterior and posterior supranasals, 2 slightly enlarged postnasal, and 1 or 2 granular scales); preorbital scales 14���20; supralabials 9���12, grading into granular scales posteriorly; infralabials 9���12; eyes large, pupils vertical; eyelid fringe scales 51���59, those of upper eyelid slightly enlarged; outer surface of upper eyelid composed of granular scales of about one half the size of those on top of head, without enlarged tubercles; fold of skin originating from suborbital region extends posteroventrally across angle of jaw; external auditory meatus elliptical with long axis directed dorsoventrally; tympanum deeply recessed; mental triangular, bordered laterally by first infralabial on each side and posteriorly by 2���4 postmentals; postmentals bordered by 7���10 gular scales; gular region below lower jaws with enlarged tubercles; gular scales juxtaposed and granular, grading posteriorly into flat hexagonal pectoral scales and even larger ventral scales. Neck narrower than body, covered with uniform granular scales interspersed with several sharply pointed conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat flatter; dorsal body tubercles surrounded by 11���13 granular scales; dorsal tubercle rows at midbody 19���24; paravertebral tubercles between limb insertions 31���37, distinct vertebral row of tubercles absent; scales around midbody 118���130; ventral scales large; males with 25���30 precloacal pores in continuous series, females without precloacal pores (but pitted scales present); region posterior to vent covered by flat juxtaposed scales and greatly swollen, containing 1 or 2 enlarged tubercles on each side at level of vent; tail thick at base; light band on tail base in width of 9���12 granular scales and with 8���10 tubercles in transversal series. Limbs relatively long and thin, covered dorsally with granular scales interspersed with several tubercles and ventrally with flat juxtaposed to subimbricate scales; dorsal granular scales grade into slightly flattened subimbricate scales on top of pes and manus; hind limbs larger than forelimbs; axillary pockets deep; subdigital lamellae wide, 10 or 11 under first finger, 18���21 under fourth finger, 11 or 12 under first toe, 21���25 under fourth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth (modified after Orlov et al. 2008; Nguyen 2011). Coloration in life (Fig. 3) Dorsal ground color of head, body and limbs signal brown (juveniles), grey-brown (young adults, SVL: 97.2���116 mm) and signal brown (adults), without small dark brown blotches (dark blotches present only on lower zone of flanks); dull white tubercles on dorsal body, limbs in juveniles, few orange tubercles on limbs in adults; iris orange or red brown; five bands on the dorsal ground, thin, immaculate without dark spots, slight bisque in juveniles, orange brown or yellow in adults, all edged anteriorly and posteriorly by thin dark brown bands, including one thin nuchal loop extending from posterior corners of eyes and posteriorly protracted (in V-shape), three body bands between limb insertions, and another one on tail base; ground color of tail dark brown, and signal brown at mottled tail base; 3���6 immaculate white caudal bands; ventral surfaces of head, body and limbs dull white and immaculate except for few dark brown spots on margin regions of gular and limbs., Published as part of Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, pp. 38-67 in European Journal of Taxonomy 751 (1) on pages 47-49, DOI: 10.5852/ejt.2021.751.1379, http://zenodo.org/record/4890455, {"references":["Orlov N. L., Ryabow S. A., Nguyen T. T., Nguyen Q. T. & Ho T. C. 2008. A new species of Goniurosaurus (Sauria: Gekkota: Eublepharidae) from north Vietnam. Russian Journal of Herpetology 15: 229 - 244.","Nguyen T. Q. 2011. Systematics, Ecology, and Conservation of the Lizard Fauna in Northeastern Vietnam, with Special Focus on Pseudocalotes (Agamidae), Goniurosaurus (Eublepharidae), Sphenomorphus and Tropidophorus (Scincidae) from this Country. PhD thesis, University of Bonn, Germany."]}

Cyrtodactylus ngati sp. nov. (Figs. 2���3) Holotype. HNUE-R00111 (Field number PT.2017.215), adult male, collected on 17 August 2017 by D. T. Le, D. T. Truong, H.Q. Nguyen, N.H. Nguyen, and A.N. Nguyen, in the karst forest near Pa Thom Cave, Pa Xa Lao Village, Pa Thom Commune, Dien Bien District, Dien Bien Province, Vietnam (21��17���N; 102��54���E, 695 m a.s.l.). Paratypes. IEBR 4829 (Field number PT.2016.07), adult female, VNUF R.2020.12 (Field number PT.2016.06), adult female, collected on 27 August 2016; HNUE-R00112 (Field number PT.2017.187), subadult female, collected on August 2017, the same data as the holotype. Diagnosis. The new Cyrtodactylus species can be distinguished from remaining congeners by the following combination of morphological characters: maximum SVL 69.3 mm; dorsal pattern consisting of six dark irregular transverse bands between limb insertions; intersupranasal single; dorsal tubercles present on occiput, body, hind limbs and on first half of tail; 17���22 irregular dorsal tubercle rows at midbody; lateral folds clearly defined, with interspersed tubercles; 32���38 ventral scales between ventrolateral folds; 13 precloacal pores separated by a diastema of 5/5 poreless scales from 7/7 femoral pores in enlarged femoral scales; precloacal and femoral pores absent in females; 1���3 postcloacal tubercles on each side; median subcaudal scales not transversely enlarged. Description of holotype. Adult male, SVL 66.5 mm; body elongate (TrunkL/SVL 0.43); head elongate (HL/ SVL 0.30), relatively wide (HW/HL 0.62), depressed (HH/HL 0.36), distinguished from neck; loreal region concave; snout long (SE/HL 0.37), obtuse anteriorly, longer than diameter of orbit (OD/SE 0.50); scales on snout small, round or oval, granular, bigger than those in frontal and parietal regions; orbit large (OD/HL 0.19), pupils vertical; supraciliaries with spinous scales posteriorly; ear oval-shaped, small (EarL/HL 0.03); rostral wider than high with a straight medial suture, bordered with first supralabial and nostril on each side; supranasals in contact with each other; nostril opening oval, surrounded by supranasal, rostral, first supralabial, and two enlarged postnasals; mental triangular, slightly wider than long (ML/MW 0.68); two enlarged postmentals; 10/9 supralabials; 8/9 infralabials; dorsal scales granular; dorsal tubercles round, conical, present from occipital region to dorsum and half of tail, tubercles in 18 irregular rows at midbody, larger at midbody, each surrounded by nine granular scales; ventrolateral body folds well defined, with interspersed tubercles; ventral scales smooth, round, midventral scales three times larger than the granules on dorsum, 38 longitudinal ventral scale rows at midbody; gular region with homogeneous, small and smooth scales; 168 ventral scales from mental to cloacal slit; precloacal groove absent; enlarged precloacal scales present; 13 contiguous precloacal pores, separated by 5/5 poreless scales and 7/7 femoral pores (FPl+PP+FPr: 7+13+7); enlarged precloacal and femoral scales present. Fore- and hind limbs moderately slender (ForeL/SVL 0.13, CrusL/SVL 0.16); enlarged tubercles on dorsal surface of forelimbs absent, but covered with slightly enlarged granules; hind limbs with distinctly developed tubercles dorsally; fingers and toes basally webbed; 16/16 lamellae under fourth finger; 12/13 lamellae under fourth toe. Tail longer than SVL (TaL 74.1 mm); 3/2 postcloacal tubercles; dorsal surface of tail bearing distinct tubercles on anterior one-half of tail; lateral rows of spinose scales present; median subcaudal scales slightly enlarged, flat, smooth. ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page Coloration in life. Background coloration light yellowish brown, with dark brown dorsal pattern; dorsal surface of head with irregular dark brown blotches; an irregularly shaped dark nuchal loop present, edged in dark brown, butterfly-shaped; six dark transversal bands between limb insertions, somewhat irregularly shaped, somewhat muted in color; tubercles at midbody greyish brown; dorsal surface of limbs with dark spots; dorsal surface of original tail dark grey with light bands, and dark narrow ring within the latter; ventral surface of head, body and limbs cream; subcaudal region dark grey with light bands (see Fig. 2). Sexual dimorphism and variation. The females differ from males by the absence of hemipenial swellings at the tail base as well as femoral and precloacal pores. For further morphological characters of the paratypes see Table 2 and Fig. 4. Comparisons. We compared Cyrtodactylus ngati sp. nov. with other congeners from Vietnam and neighboring countries in the mainland Indochina region, including Laos, Cambodia, and Thailand based on examination of specimens (see Appendix) and data from literature (Luu et al. 2014; Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014 a,b; Nurngsomsri et al. 2014; Grismer et al. 2015; Luu et al. 2015; Sumontha et al. 2015; Pauwels et al. 2016; Le et al. 2016; Luu et al. 2016a,b,c, 2017; Nguyen et al. 2017; Chuaynkern et al. 2018; Pauwels et al. 2018; Nazarov et al. 2018; Murdoch et al. 2019; Pham et al. 2019; Sitthivong et al. 2019; Schneider et al. 2020; Ostrowski et al. 2020, 2021). The new species can be differentiated from other known species of the genus Cyrtodactylus by morphological characters (see Table 3). Below we compared the new species with the Cyrtodactylus brevipalmatus group data obtained from Smith (1923); Dring (1979); Ulber (1993); Nurngsomsri et al. (2014); and Chuaynkern et al. (2018). In comparison with the species from the brevipalmatus group, comprising C. interdigitalis Ulber, C. elok Dring, and C. brevipalmatus (Smith), the new species can be distinguished from C. interdigitalis by its smaller size (maximum SVL 69.3 mm versus 80.0 mm), having fewer precloacal pores (13 versus 14���16), more body bands (6 versus 3), the absence of tubercles on forelimbs (versus present), the presence of dark narrow ring in each light tail band (versus absent), and the presence of slightly enlarged subcaudal scales (versus absent); from C. elok by its smaller size (maximum SVL 69.3 mm versus 83.0 mm), having fewer ventral scales (32���38 versus 52), more body bands (6 versus 3), more precloacal pores in males (13 versus 8), the presence of enlarged femoral scales and femoral pores in males (versus absent), and the presence of slightly enlarged subcaudal scales (versus absent); from C. brevipalmatus by having more femoral pores in males (13 versus 9 or 10), more body bands (6 versus 3), the absence of precloacal pores in females (versus present), and dorsal pattern banded (versus blotched). For more details see Table 4. Distribution. Cyrtodactylus ngati sp. nov. is currently known only from the type locality in Dien Bien Province, Vietnam (Fig. 4). Etymology. We name this new species in honor of our colleague, Assoc. Prof. Dr. Ngat Nguyen Le, Hanoi National University of Education, Vietnam, in recognition of his contribution to herpetological research in Vietnam. As common names, we suggest Ngat���s Bent-toed Gecko (English) and Th���n l���n ng��n ng���t (Vietnamese). Ecological notes. Specimens were found between 19:00 and 21:00, on limestone cliffs and in rock crevices, approximately 1���2 m above the ground. The surrounding habitat was disturbed evergreen forest of small hardwoods and shrubs on the mountain slope. The humidity was approximately 74���76% and the air temperature ranged from 23 to 26 oC (Fig. 5). All other members of the Cyrtodactylus brevipalmatus group are arboreal, have prehensile tails carried above their back in a coiled position, and usually are found only on vegetation (Grismer et al. 2020). This is the first member of the group that apparently lacks a prehensile tail and occurs on karst. Grismer et al. (2020) demonstrated that an arboreal habitat preference was the ancestral condition for the C. brevipalmatus group and these findings represent tthe first record of a potential transition of an arboreal habitat to a karst habitat preference in Cyrtodactylus., Published as part of Le, Dzung Trung, Sitthivong, Saly, Tran, Tung Thanh, Grismer, L. Lee, Nguyen, Truong Quang, Le, Minh Duc, Ziegler, Thomas & Luu, Vinh Quang, 2021, First record of the Cyrtodactylus brevipalmatus group (Squamata: Gekkonidae) from Vietnam with description of a new species, pp. 492-510 in Zootaxa 4969 (3) on pages 495-505, DOI: 10.11646/zootaxa.4969.3.3, http://zenodo.org/record/4751410, {"references":["Luu, Q. V., Calame, T., Bonkowski, M., Nguyen, Q. T. & Ziegler, T. (2014) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Khammouane Province, Laos. Zootaxa, 3760 (1), 54 - 66. https: // doi. org / 10.11646 / zootaxa. 3760.1.3","Nazarov, R. A., Poyarkov, N. A., Orlov, N. L., Nguyen, S. N., Milto, K. D., Martynov, A. A., Konstantinov, E. L. & Chulisov, A. S. (2014) A review of genus Cyrtodactylus (Reptilia: Sauria: Gekkonidae) in fauna of Laos with description of four new species. Proceedings of the Zoological Institute RAS, 318, 391 - 423.","Nguyen, S. N., Yang, J. - X., Le, T. - N. T., Nguyen, L. T., Orlov, N. L., Hoang, C. V., Nguyen, T. Q., Jin, J. - Q., Rao, D. - Q., Hoang, T. N., Che, J., Murphy, R. W. & Zhang, Y. - P. (2014) DNA barcoding of Vietnamese bent-toed geckos (Squamata: Gekkonidae: Cyrtodactylus) and the description of a new species. Zootaxa, 3784 (1), 48 - 66. https: // doi. org / 10.11646 / zootaxa. 3784.1.2","Panitvong, N., Sumontha, M., Tunprasert, J. & Pauwels, O. S. G. (2014) Cyrtodactylus saiyok sp. nov., a new dry evergreen forest-dwelling Bent-toed Gecko (Squamata: Gekkonidae) from Kanchanaburi Province, western Thailand. Zootaxa, 3869 (1), 64 - 74. https: // doi. org / 10.11646 / zootaxa. 3869.1.6","Schneider, N., Nguyen, T. Q., Le, M. D, Nophaseud, L., Bonkowski, M. & Ziegler, T. (2014 a) A new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of northern Laos. Zootaxa, 3835 (1), 80 - 96. https: // doi. org / 10.11646 / zootaxa. 3835.1.4","Nurngsomsri, P., Chuaynkern, Y., Chuaynkern, C., Duengkae, P., Sribandit, P., Wongnak, T., Ittiporn, K., Ouchan, C. & Eimampai, K. (2014) Variation and recent distribution of Cyrtodactylus interdigitalis Ulber, 1993 from Thailand. In: Proceedings of the 40 th Congress on Science and Technology of Thailand (STT 40). Khon Kaen University, Khon Kaen, 2 December 2014, pp. 753 - 759.","Grismer, L. L., Wood, Jr., P. L., Ngo, V. T. & Murdoch, M. L. (2015) The systematics and independent evolution of cave ecomorphology in distantly related clades of Bent-toed Geckos (Genus Cyrtodactylus Gray, 1827) from the Mekong Delta and islands in the Gulf of Thailand. Zootaxa, 3980 (1), 106 - 126. https: // doi. org / 10.11646 / zootaxa. 3980.1.6","Sumontha, M., Pauwels, O. S. G., Panitvong, N., Kunya, K. & Grismer, J. L. (2015) A new lowland forest Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Ranong Province, peninsular Thailand. Zootaxa, 3911 (1), 106 - 118. https: // doi. org / 10.11646 / zootaxa. 3911.1.6","Luu, V. Q., Calame, T., Nguyen, T. Q., Bonkowski, M. & Ziegler, T. (2015) A new species of Cyrtodactylus (Squamata: Gekkonidae) from limestone forest, Khammouane Province, central Laos. Zootaxa, 4058 (3), 388 - 402. https: // doi. org / 10.11646 / zootaxa. 4058.3.6","Luu, V. Q., Bonkowski, M., Nguyen, T. Q., Le, M. D., Schneider, N., Ngo, H. T. & Ziegler, T. (2016 a) Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos. Zootaxa, 4107 (2), 101 - 140. https: // doi. org / 10.11646 / zootaxa. 4107.2.1","Le, D. T., Nguyen, Q. T., Le, M. D. & Ziegler, T. (2016) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Ninh Binh Province, Vietnam. Zootaxa, 4162 (2), 268 - 282. https: // doi. org / 10.11646 / zootaxa. 4162.2.4","Luu, V. Q., Dung, T. V., Nguyen, T. Q., Le, M. D. & Ziegler, T. (2017) A new species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from Gia Lai Province, Central Highlands. Zootaxa, 4362 (3), 385 - 404. https: // doi. org / 10.11646 / zootaxa. 4362.3.4","Nguyen, T. Q., Pham, A. V., Ziegler, T., Ngo, H. T. & Le, M. D. (2017) A new species of Cyrtodactylus (Squamata: Gekkonidae) and the first record of C. otai from Son La Province, Vietnam. Zootaxa, 4341 (1), 25 - 40. https: // doi. org / 10.11646 / zootaxa. 4341.1.2","Pauwels, O. S. G., Nazarov, R. A., Bobrov, V. V. & Poyarkov, N. A. (2018) Taxonomic Status of two populations of Bent-toed Geckos of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) with description of a new species from Nui Chua National Park, southern Vietnam. Zootaxa, 4403 (2), 307 - 335. https: // doi. org / 10.11646 / zootaxa. 4403.2.5","Chuaynkern, Y., Nurngsomsri, P., Chuaynkern, C., Duengkae, P. & Karaphan, S. (2018) Short Communication: Cyrtodactylus elok Dring, 1979 (Sauria, Gekkonidae): A first country record for Thailand. Biodiversitas, 19, 2111 - 2117. https: // doi. org / 10.13057 / biodiv / d 190617","Nazarov, R. A., Pauwels, O. S. G., Konstantinov, E. L., Chulisov, A. S., Orlov, N. L. & Poyarkov, N. A. (2018) A new Karst-dwelling bent-toed gecko (Squamata: Gekkonidae: Cyrtodactylus) from Xiangkhoang Province, northeastern Laos. Zoological Research, 39 (3), 197 - 213.","Murdoch, M. L., Grismer, L. L., Wood JR, P. L., Neang, T., Poyarkov, N. A., Ngo, T. V., Nazarov, R. A., Aowphol, A., Pauwels, O. S. G., Nguyen, H. N. & Grismer, J. L. (2019) Six new species of the Cyrtodactylus intermedius complex (Squamata: Gekkonidae) from the Cardamom Mountains and associated highlands of Southeast Asia. Zootaxa, 4554 (1), 1 - 62. https: // doi. org / 10.11646 / zootaxa. 4554.1.1","Pham, A. V., Le, M. D., Ngo, H. T., Ziegler, T. & Nguyen, T. Q. (2019) A new species of Cyrtodactylus (Squamata: Gekkonidae) from northwestern Vietnam. Zootaxa, 4544 (1), 360 - 380. https: // doi. org / 10.11646 / zootaxa. 4544.3.3","Sitthivong, S., Luu, V. Q., Ha, N. V., Nguyen, T. Q., Le, M. D. & Ziegler, T. (2019) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Vientiane Province, northern Laos. Zootaxa, 4701 (3), 257 - 275. https: // doi. org / 10.11646 / zootaxa. 4701.3.3","Schneider, N., Luu, V. Q., Sitthivong, S., Teynie, A., Le, M. D., Nguyen, T. Q. & Ziegler, T. (2020) Two new species of Cyrtodactylus (Squamata: Gekkonidae) from northern Laos including new finding and expanded diagnosis of C. bansocensis. Zootaxa, 4822 (4), 503 - 530. https: // doi. org / 10.11646 / zootaxa. 4822.4.3","Ostrowski, S., Do, D. T, Le, M. D., Ngo, H. T., Pham, C. T., Nguyen, T. Q., Nguyen, V. T. & Ziegler, T. (2020) A new species of Cyrtodactylus (Squamata: Gekkonidae) from southern Vietnam. Zootaxa, 4789 (1), 171 - 203. https: // doi. org / 10.11646 / zootaxa. 4789.1.5","Ostrowski, S., Le, M. D., Ngo, H. T., Pham, C. T., Phung, T. M, Nguyen, T. Q. & Ziegler, T. (2021) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Binh Thuan Province, southern Vietnam. European Journal of Toxonomy, 731, 47 - 70. https: // doi. org / 10.5852 / ejt. 2021.731.1203","Pauwels, O. S. G., Sumontha, M. & Bauer, A. M. (2016) A new Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Phetchaburi Province, Thailand. Zootaxa, 4088 (3), 409 - 419. https: // doi. org / 10.11646 / zootaxa. 4088.3.6","Smith, M. A. (1923) Notes on reptiles and batrachians from Siam and Indo-China (No. 2). Journal of Natural History Society of Siam, 6, 47 - 53.","Dring, J. C. M. (1979) Amphibians and reptiles from northern Trengganu, Malaysia, with descriptions of two new geckos: Cnemaspis and Cyrtodactylus. Bulletin of the British Museum (Natural History), Zoology, 34 (5), 181 - 24.","Ulber, T. (1993) Bemerkungen uber cyrtodactyline Geckos aus Thailand nebst Beschreibungen von zwei neuen Arten (Reptilia: Gekkonidae). Mitteilungen aus dem Zoologischen Museum in Berlin, 69 (2), 187 - 200. https: // doi. org / 10.1002 / mmnz. 4840690202","Grismer, L. L., Chan, K. O., Oaks, J. R., Neang T., Sokun, L., Murdoch, M. L., Stuart, B. L. & Grismer, J. L. (2020) A new insular species of the Cyrtodactylus intermedius (Squamata: Gekkonidae) group from Cambodia with a discussion of habitat preference and ecomorphology. Zootaxa, 4830 (1), 75 - 102. https: // doi. org / 10.11646 / zootaxa. 4830.1.3"]}

Dixonius somchanhae sp. nov. Figs. 2–5 Holotype: Adult male, VNUF R.2020.3 (Field no. VT03) in Huaysorn-Huaysua Village, Nasaithong District, Vientiane Capital (18 o 11.450’ N, 104 o 45.926’ E, at an elevation of 285 m a.s.l.), Laos, collected by V. Q. Luu, N. V. Ha and S. Sitthivong on 26 September 2016. Paratypes. VNUF R.2020.1 (Field no. VT01), adult male, and VNUF R.2020.2 (Field no. VT02), adult m a le; the same data as given for the holotype. VNUF R.2020.4 (Field no. VT02-14), adult female, and VNUF R.2020.5 (Field no. VT03-14), adult female, collected by V.Q. Luu and T. Calame in June 2014. Diagnosis. A species of the genus Dixonius, characterized by a combination of the following characters: maximum SVL 47.1 mm; 19–21 longitudinal rows of dorsal tubercles at midbody; 23–26 longitudinal rows of ventrals across the abdomen; 7 or 8 supralabials, 6 in mid-orbital position; 5 or 6 infralabials; 5 or 6 precloacal pores in males, femoral pores lacking; precloacal and femoral pores absent in females; a canthal stripe running from rostrum through the eye and terminating at back of head; uniformly brown dorsum. Description of the holotype. Adult male, SVL 43.8 mm with dorsolaterally flattened body, TaL 62.7 mm, trunk length 20.5 mm. Head longer than wide, depressed, distinct from neck. Head length 12.2 mm, head width 8.5 mm, eye of moderate size 3.4 mm, ear-opening round (EL 1.6 mm), snout-eye length 5.1 mm, internarial distance 1.3 mm. Rostral very large, wider than high (width 1.9 mm, height 1.4 mm), with distinct suture; supralabials 7/7 (6 in midorbital position); nostril in contact with rostral, first supralabial, supranasal, and two nasals posteriorly on each side; supranasals in broad contact with each other; without intersupranasals; snout flat, covered with granular scales; pupil vertical; ear opening round, approximately one half of the eye diameter, without bordering enlarged scales; infralabials 5/5, decreasing gradually in size; mental triangular, wider than high (width 2.0 mm, height 1.6 mm); two pairs of enlarged postmentals, first pair very large and in contact with each other, second pair about one third size of first, in contact with the first and second infralabials and separated from each other by seven gular scales; dorsal tubercle rows at midbody 20, keeled, separated from each other by one smaller scale, keeled or at least almost conical shaped; ventral scales larger than dorsal scales, in 24 rows at midbody; dorsal surface of fore- and hind limbs covered by feebly keeled scales, area around limb insertions covered with small granular scales; paravertebral scales (number of scales in a paravertebral row from first scale posterior to parietal scale to last scale at the level of cloaca) 35; paravertebral scales in a row between limb insertions 21; lamellae on fourth toe 14; precloacal pores 6, in an angular series; tail covered by keeled scales, in different sizes; subcaudals 56, enlarged and undivided. Coloration in life. Dorsum of the holotype brown in life with irregular dark marks (Fig. 2); head with dark blotches; a light brown canthal stripe running from rostrum through eyes, ending at back of head; light spots irregularly arranged from head to tail; ventral surface light beige to uniformly whitish as the belly and the throat; supralabials grey with dark grey spots; upper surface of toes uniformly light grey with dark blotches; dorsal surface of the tail light brownish-grey with cream lines. In preservative, dorsal head, back and dorsal surface of tail brownish grey with irregular dark brown blotches; coloration on dorsal surface of fore- and hindlimbs grey brown. Canthal stripe dark brown, running from rostrum through eyes, stretching behind orbit to back of head. Round white spots arranged in two irregular rows extend from back of head along body and tail. Sexual dimorphism. Morphologically, paratypes corresponded well with the holotype, except for the absence of precloacal pores in females (Table 2). Comparisons. The new species from Vientiane Capital, Laos can be distinguished from all known Dixonius species as follows: from D. taoi from Vietnam by having more dorsal tubercle rows (19–21 versus 11–12), more ventral scale rows at midbody (23–26 versus 21–23), and different color pattern on head and tail (light spots irregular arranged from head to tail versus four to seven round yellowish spots, arranged in one or two irregular rows from back of head onwards and continuing along tail); from D. minhlei from Vietnam by having more dorsal tubercle rows (19–21 versus 14–15), fewer precloacal pores in males (5–6 versus 7–8), and more ventral scale rows at midbody (23–26 versus 20–23); from D. siamensis by its smaller size (maximum SVL 47.1 versus 57 mm), having more dorsal tubercle rows (19–21 versus 10–14), and generally fewer precloacal pores in males (5–6 versus 6–7); from D. aaronbaueri by having more ventral scale rows at midbody (23–26 versus 18–19), more dorsal tubercle rows (19–21 versus 11), and fewer paravertebral scales in a row between limb insertions (PV’ 19–27 versus 29– 32); from D. dulayaphitakorum by having more ventral scale rows at mid-body (23–26 versus 22), fewer dorsal tubercle rows (19–21 versus 22), fewer interorbital scale (7–8 versus 9–10), and fewer infralabials (5–6 versus 7); from D. pawangkhananti by having more ventral scale rows at mid-body (23–26 versus 16), more dorsal tubercle rows (19–21 versus 16), and more paravertebral scales from first scale posterior to parietal scale to last scale at the level of cloaca (PV 35–40 versus 30–32); from D. hangseesom by having more dorsal tubercle rows at midbody (19–21 versus 12–14), more paravertebral scales from first scale posterior to parietal scale to last scale at the level of cloaca (PV 35–40 versus 26), and fewer precloacal pores in males (5–6 versus 7–8); from D. melanostictus by having more ventral scale rows at midbody (23–26 versus 22), more dorsal tubercle rows (19–21 versus 10–11), more paravertebral scales in a row between limb insertions (PV’ 19–27 versus 10–11), and fewer precloacal pores in males (5–6 versus 9); from D. kaweesaki by having more dorsal tubercle rows (19–21 versus 12–13), fewer precloacal pores in males (5–6 versus 9–11), different canthal stripe (a light brown canthal stripe running from rostrum through eyes, ending at back of head versus a black canthal stripe running through the eye, joining the upper part of the flanks), and coloration of tail light brownish-grey (versus light orange); from D. vietnamensis by its larger size (maximum SVL 47.1 versus 40.8 mm), having more dorsal tubercle rows at midbody (19–21 versus 16), and more supralabials (7–8 versus 5–6); from D. lao by its smaller size (maximum SVL 47.1 versus 55.4 mm), fewer infralabials (5–6 versus 7–8), and fewer precloacal pores in males (5–6 versus 8). For further distinguishing characters see Table 3. Distribution. Dixonius somchanhae sp. nov. is currently known only from the type locality in Huaysorn- Huaysua Village, Vientiane Capital, Laos (Fig. 6). Natural history notes. The type series was found at night, between 19h00 and 22h00, on the rocks of sandstone area (Fig. 7). The surrounding habitat was disturbed secondary forest of small hardwoods and shrubs. Etymology. The new Dixonius somchanhae species is named after Associate Professor Dr. Somchanh Bounphanmy, from the National University of Laos, in recognition of her support of our biodiversity research in Laos.

Cyrtodactylus chungi sp. nov. urn:lsid:zoobank.org:act: C3B181E8-F9D3-4344-AD20-28765D1BA0B2 Figs 3–8 Diagnosis The new species can be distinguished from other members of the genus Cyrtodactylus by a combination of the following characters (Table 1): relatively small body size (SVL up to 68.5 mm); a continuous neckband; 5 or 6 irregular transverse dorsal bands; 11 or 12 bands on original tail; keeled tubercles present on dorsum, posterior limbs and tail; 17 or 18 irregular dorsal tubercle rows; 30 or 31 ventral scale rows; ventrolateral skin folds indistinct; an angular series of seven precloacal pores in male and six pitted, enlarged precloacal scales in female, each series separated by a diastema of undifferentiated scales from 4–6 enlarged, poreless femoral scales; median subcaudals slightly enlarged; 17–20 subdigital lamellae under the fourth toe. Etymology Specific epithet ‘ chungi ’ is in honor of Prof. Dr Ngo Dac Chung, College of Education, Hue University, in recognition of his outstanding contributions to training of many herpetologist generations from Hue University. For the common names we suggest Chung’s Bent-toed Gecko (English) and Thạch sùng ngón chứng (Vietnamese). Type material (Figs 3–8) Holotype VIETNAM • ♂; Southern Vietnam, Ham Thuan Nam District, Binh Thuan Province, Ta Kou Nature Reserve; 10 o 48.868′ N, 107 o 53.523′ E; 424 m a.s.l.; 20 Jun. 2017; T.M. Phung leg.; Field No. BT2017.1; sequence with GenBank accession number MT576019; IEBR 4581. Paratype VIETNAM • 1 ♀; same collection data as for holotype; Field No. BT2017.2; sequence with GenBank accession number MT576020; IEBR 4582. Description of holotype Adult male, snout-vent length (SVL) 66.6 mm; tail length (TL) 62.7 mm (45.1 mm regenerated); head distinct from neck, relatively long (HL: 20.1 mm; HL/SVL ratio 0.32), HW 13.3 mm (HW/ HL ratio 0.66), HH 7.8 mm (HH/HL ratio 0.4); loreal region slightly bulged; snout elongated and rounded (SE: 7.6; SE/HL ratio 0.38); eye diameter 4.4 mm, more than half of snout-eye distance (OD/SE ratio 0.58); median snout scales small and rounded; eyes relatively small compared to head (OD/HL ratio 0.22); anterior supraciliaries large and rounded, posterior supraciliaries small and tapered; ear openings relatively small (ED: 0.75 mm; ED/HL ratio 0.04), vertical and oval in shape; eye to ear distance slightly longer than eye diameter (EyeEar/ED ratio 1.39); rostral wider than high (RH: 1.65 mm; RW: 3.3 mm; RW/RH ratio 2), with an inverse, Y-shaped median suture, surrounded by first supralabial, two nostrils, two supranasals and internasal; lateral nostrils elongated, oval and surrounded by rostral, first supralabial, three postnasals and one supranasal; supranasals separated by one internasal; eye separated from supralabials by 3 or 4 scale rows; mental triangular, wider than high (MW: 1.75 mm; MH 2.6 mm; MW/MH ratio 1.49), surrounded by first two infralabials and posteriomedially by postmentals; supralabials 10/9; infralabials 9/9; body slender and short (AG/SVL ratio 0.41); ventrolateral skin folds not developed; dorsal scales granular, heterogeneous in shape and size; dorsal tubercles present in posterior part of head, extremities, dorsum and first third of tail; postocular tubercles irregularly distributed, rounded, flat and about three times size of surrounding granular scales; median dorsal tubercles about five times as large as granular scales, oval, and not keeled; posterior dorsal tubercles keeled and on sacral region very strongly keeled up to base of tail; 18 irregular transversal dorsal tubercle rows; 24 paravertebral tubercles; each tubercle surrounded by 9 or 10 granular scales; ventrals smooth, oval and slightly imbricate, in 31 longitudinal rows; gular scales homogeneous; dorsal surfaces of upper and lower arms without tubercles, but occasionally enlarged granular scales; left thigh with series of 6 enlarged femoral scales separated by diastema of 6 smaller scales from precloacal scales; right thigh with series of 3 enlarged femoral scales separated by diastema of 8 smaller scales from precloacal scales and single enlarged femoral scale distally; no femoral pores; 45 enlarged precloacal scales arranged in diamond shape, with angled series of 7 pore-bearing scales (3 right, 1 angled, 3 left); 15 posteriorly enlarged scales and 21 anteriorly enlarged scales; two postcloacal tubercles on each side; scales on palms and soles rounded to oval, smooth and arched; fore- and hindlimbs moderately long (ForeaL/SVL ratio 0.15; Tibia/SVL ratio 0.17) relatively narrow; toes and fingers narrow and curved at interphalangeal joint; slightly transparent, pointed and curved claws, bordered ventrally by large lamella and dorsally by small, narrow scale; subdigital lamellae proximally larger, widened and merge basally into smaller scales of palm and sole surfaces; distal lamellae smaller, more elongated and distally increasingly imbricate; right fourth finger with 15 subdigital lamellae (6 proximal, 9 distal); left fourth finger with 16 subdigital lamellae (5 proximal, 11 distal); right and left fourth toe with 17 subdigital lamellae each (6 proximal, 11 distal); base of tail with small granular supracaudals and strongly keeled, rounded tubercles; last two thirds of tail regenerated, scales heterogeneously arranged and shaped; tubercles of original part arranged in 6 transverse rows (8/-/4/4/4/4) in a distance of 6–7 granular scale rows; tubercles slightly keeled; subcaudals of original part oval, rounded and imbricate; median subcaudals about two to three times larger than the lateral scales, but not transversely enlarged, or arranged in definable rows. Coloration in preservative (Figs 5–8) Ground color of head, neck, limbs and tail grey to light brown; dorsal surface of head brown with irregular, small, dark brown spots; two larger dark brown blotches located in posterior interorbital area and two other dark brown blotches in occiput area; skin above eyes dark grey, blue to black; rostral dark brown; supralabials and infralabials also brown, turning to cream at adjacent areas; a distinct dark brown stripe in loreal region, reaching to third supralabial; nuchal loop dark brown, in V-shape, extending from posterior corner of eye to the neck, edged in light grey; tubercles on head, limbs, dorsum and tail light to dark brown; tubercles on lateral skin fold white to gray; dorsal surfaces of limbs with small, irregularly distributed, dark brown spots; dorsal surface of toes and fingers dark brown spotted with light grey; dorsum with 5 irregular transverse dark brown bands; first band over the shoulder region, X-shaped; dark and light brown spots present between transverse bands and on flanks; ventral scales of body light grey to cream; lateral sides of head, below postocular stripe, and ventrolateral margins of flanks slightly white to cream; anterior part of tail with two transverse, dark brown bands dorsally with dark brown spots in interspaces; dark, transverse dorsal bands separated from underside of tail by light brown, whitish spotted sides of tail; tail light grey to cream ventrally and marbled brown on the original part; regenerated part of tail grey to light brown, marbled. Coloration in life Ground color of dorsal surface of head, limbs and dorsum bright-yellow; transverse body and tail bands darker; lateral head and belly with bright-yellow spots; marbling of surface of limbs darker; ventral surface white to light blue (Figs 3–4). Sexual dimorphism and variation The morphological characteristics of the female IEBR 4582 correspond well with the holotype (IEBR 4581) and differ only in a few details. The female has some slightly keeled tubercles on the dorsal surface of the forelimbs. Dorsal tubercles are keeled from the posterior insertion of the forelimbs. The two supranasals are separated by two internasals. Another internasal is inserted between the rostral and the two internasals. Two inverse sutures are X-shaped from the lower edges of the internasal and do not form a median suture. The tail is original. The median subcaudals are about twice as large as the supracaudals and extend to the tip of the tail. The two dark brown spots of the occiput are longer and larger. The neck band is also V-shaped but pointed posteriorly. The dorsal surfaces of the forearms and hindlimbs with fine, irregular, transverse stripes. Dorsum with 6 transverse, irregular bands. The first band, between the insertion of the forelimbs, is clearly X-shaped over the shoulder region. Between the neck band and the first body band is an oval, dark brown spot. The dorsal tubercles are clearly lighter. The dorsal side of the tail bears 11–12 transverse, dark brown bands, which completely enclose only the last third. Between these bands are irregular, fine, light brown to brown stripes and dots. The first two thirds of the underside of the tail are marbled light grey to brown. The last third resembles the dorsal side. Comparison We compared the new species with its 19 congeners from the Cyrtodactylus irregularis complex based on an examination of specimens (Table 2) and data obtained from the literature (Smith 1921 a, 1921b; Ziegler et al. 2002, 2013; Heidrich et al. 2007; Orlov et al. 2007; Nazarov et al. 2008. 2012; Ngo & Bauer 2008; Rösler et al. 2008; Geissler et al. 2009; Ngo & Chan 2010; Ngo 2013; Nguyen et al. 2013; Schneider et al. 2014; Luu et al. 2017; Pauwels et al. 2018; Neang et al. 2020; Ostrowski et al. 2020). The new species can be distinguished from all other Cyrtodactylus species from Vietnam by morphological characteristics (see Table 3). In comparison with the species from the C. irregularis complex Cyrtodactylus chungi sp. nov. is distinguished from C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen, 2007, C. kingsadai Ziegler, Phung, Le & Nguyen, 2013, C. takouensis Ngo & Bauer, 2008 and C. yangbayensis Ngo & Chan, 2010 by the absence of transversely enlarged subcaudals. The new species can be distinguished from C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler, 2007, C. gialaiensis Luu, Tran, Nguyen, Le & Ziegler, 2017, C. pseudoquadrivirgatus Rösler, Nguyen, Vu, Ngo & Ziegler 2008 and C. taynguyenensis Nguyen, Le, Tran, Orlov, Lathrop Macculoch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, 2013 by the presence of enlarged femoral scales. The new species can be distinguished from C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, 2012 by its smaller size (SVL of 66.6–68.5 vs 74.0– 86.3 mm), fewer ventral scale rows (30–31 vs 38–43) and enlarged femoral scales (4–6 vs 8–10), more precloacal pores in the male (7 vs 4–6), the presence of 6 pitted precloacal scales in the female (vs absence) and by the different dorsal color patter (irregular transversal bands vs irregular transversal bands with light borders). Cyrtodactylus chungi sp. nov. differs from C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, 2012 by having fewer ventral scale rows (30–31 vs 36–46) and enlarged femoral scales (4–6 vs 6–8), and by the different dorsal color pattern (irregular transversal bands vs round spots forming irregular transverse bands). Cyrtodactylus chungi sp. nov. differs from C. cucdongensis Schneider, Phung, Le, Nguyen & Ziegler, 2014 by its larger size (SVL 66.6–68.5 vs 55.8–65.9 mm), fewer ventral scale rows (30–31 vs 35–44) and enlarged femoral scales (4–6 vs 5–9). Cyrtodactylus chungi sp. nov. differs from C. dati Ngo, 2013 by having fewer ventral scale rows (30–31 vs 42–48) and by the different dorsal color pattern (irregular transverse bands vs irregular dark blotches). Cyrtodactylus chungi sp. nov. differs from C. huynhi Ngo & Bauer, 2008 by having fewer ventral scale rows (30–31 vs 43–46) and by the absence of femoral pores (vs 3–8). Cyrtodactylus chungi sp. nov. differs from C. irregularis (Smith, 1921) by its smaller size (SVL of 66.6–68.5 vs 72.0–86.0 mm), having fewer ventral scale rows (30–31 vs 38–45) and enlarged femoral scales (4–6 vs 7–8), and by the different dorsal color pattern (irregular transverse bands vs transverse bands with uneven margins in white binding, some bands can fall into separated spots). Cyrtodactylus chungi sp. nov. differs from C. phnomchiensis Neang, Henson & Stuart, 2020 by its smaller size (SVL 66.6–68.5 vs 76.1–80.7 mm), having fewer ventral scale rows (30–31 vs 45–54) enlarged femoral scales (4–6 vs 0–8), precloacal pores in males (7 vs 4–5) and subdigital lamellae under the fourth to (17–20 vs 20–23) and by the different dorsal color pattern (irregular transverse brown bands vs dark brown body bands bordered by yellowish white or light brown bands). Cyrtodactylus chungi sp. nov. differs from C. phumyensis Ostrowski, Do, Lee, Ngo, Pham, Nguyen, Nguyen & Ziegler, 2020 by having fewer ventral scale rows (30–31 vs 33–40), the absence of pitted precloacal scales in the male (absent vs 1), more paravertebral tubercles (24–27 vs 20–23) and by the different dorsal color pattern (irregular transverse bands vs anteriorly irregularly spotted and posteriorly banded). Cyrtodactylus chungi sp. nov. differs from C. phuocbinhensis Nguyen, Le, Tran, Orlov, Lathrop Macculoch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, 2013 by its larger size (SVL 66.6–68.5 mm vs 46.0– 60.4 mm), having fewer ventral scale rows (30–31 vs 43–47), the presence of 6 pitted precloacal scales in the female (vs absence) and the different dorsal color pattern (irregular transversal bands vs stripes). Cyrtodactylus chungi sp. nov. differs from C. ziegleri Nazarov, Orlov, Nguyen & Ho, 2008 by its smaller size (SVL of 66.6–68.5 vs 84.6–93.0 mm), having fewer ventral scale rows (30–31 vs 33–39) and enlarged femoral scales (4–6 vs 8–10). Cyrtodactylus chungi sp. nov. is most similar to C. sangi Pauwels, Nazarov, Bobrov & Poyarkov, 2018. However, the new species can be distinguished from C. sangi by its larger size (SVL 66.6–68.5 vs 49.9–56.3 mm), having fewer ventral scale rows (30–31 vs 37) and by the different dorsal color pattern (irregular transversal bands with a closed neck band vs irregular transversal bands and pattern with an interrupted neck band). Distribution Cyrtodactylus chungi sp. nov. is currently known only from the type locality in Ta Kou Nature Reserve, Binh Thuan Province, Vietnam (Fig. 1). Natural history Specimens were found at night between 20:00 and 22:00, on granite rock nearby a forest path, about 0.5–1.0 m above the ground, at elevations between 400 and 500 m a.s.l. The surrounding habitat was evergreen forest of medium and small hardwoods mixed with shrubs and vines. The humidity was approximately 68–72% and the air temperature ranged from 26.8 to 31.6 oC (Fig. 9).

Ostrowski, Sabrina, Ngo, Hanh Thi, Pham, Cuong The, Phung, Trung My, Nguyen, Truong Quang, Ziegler, Thomas (2021): A new Cyrtodactylus (Squamata: Gekkonidae) from Binh Thuan Province, southern Vietnam. European Journal of Taxonomy 731: 47-70, DOI: https://doi.org/10.5852/ejt.2021.731.1203

Janssen, Helen Yvonne, Jin-Long, Ren, Jia-Tang, Li, Zeng, Wang, Nguyen, Tao Thien, Nguyen, Truong Quang, Bui, Quynh Thi Thuy, Ngo, Hanh Thi, Le, Minh Duc, Ziegler, Thomas (2020): Range extension and extended diagnosis of Lycodon pictus: First country record from China. Revue suisse de Zoologie 127 (2): 413-422, DOI: 10.35929/RSZ.0030

Lycodon pictus Janssen, Pham, Ngo, Le, Nguyen & Ziegler, 2019 Figs 1-3 New record for Vietnam (n = 1): VNMN 011227 (Fig. 1), Ha Lang, Cao Bang, Vietnam, Coordinates 22°42’50’’ N, 106°40’74’’ E, 960 m a.s.l., collected by Nguyen Quoc Huy on 18 May 2019. New record for China (n = 1): CIB 115609 (field no. GX2019069; Figs 2-3), Nonggang National Nature Reserve, Longzhou County, Guangxi Zhuang Autonomous Region, China, 106°57’18.77” E, 22°28’3.92” N, 233 m a.s.l., collected by Ren Jin-Long and Zeng Fan-Xing on 8 July 2019. Description of the Chinese specimen: Head elongate, moderately distinct from neck, rather flattened, longer than wide, snout narrow, distinctly shorter than head width; nostril large, lateral, located in the middle of the nasal; eye large, pupils vertically elliptic; rostral triangular, much broader than high, hardly visible from above; nasal divided, narrowed medially; two internasals, anteriorly round, slightly wider than high, bordered by two large, hexagonal prefrontals posteriorly; frontal single, enlarged, pentagonal to hexagonal, narrowed posteriorly; parietals large, longer than wide, in contact with each other medially, with upper anterior and posterior temporals, paraparietal laterally and four nuchal scales posteriorly; paraparietals elongated, anterior part widened, about equal in length or slightly longer than posterior temporals; loreal 1/1, elongate, not entering orbit on right side, whereas the posterior corner entering orbit on the left side (Fig. 3C and D); supralabials 8/8, first and second in contact with nasal, third to fifth entering orbit, sixth largest, slightly higher than third on left side; infralabials 10/9, first pair in broad contact with each other, first to fifth/first to fourth in contact with anterior pair of chin shields; anterior and posterior pairs of chin shields elongate, second pair not meeting in midline; preocular 1/1, located on antero-upper part of eye; postoculars 2/2, lowermost smaller, bordering anterior temporals; anterior temporal 1/1, in contact with sixth supralabial; posterior temporals 3/3, upper one thinner than lower one. Left maxilla arched, with an angular apex, distinctly bent inwards anteriorly. A total of 13 maxillary teeth or teeth alveola, with the following formula: five small anterior teeth, slightly enlarged posteriorly + three strongly enlarged teeth, thick, and not much curved + a distinctly wide gap + three small teeth + a small gap + two enlarged posterior teeth. Body elongate, TL 591 mm; SVL 467 mm; TaL 124 mm; preventral 1, ventrals 218, from behind neck region distinctly notched laterally; subcaudals 90, paired; precloacal plate single; DSR 17-17-15, all smooth; the vertebral scales not enlarged. Coloration in preservative: Dorsal surface of head and neck with pigmentation, paler on lateral side of head, the lower part of supralabials yellowish cream, speckled with pale brownish blotches. Body brownish black, light body bands beginning after 1.5 times the head length behind the head, in total 29 transverse light bands on body and 14 light bands on tail; the first four body bands yellowish cream, and distinctly widened towards the venter, increased in size posteriorly; a dark mottling in the vertebral region more prominent posteriorly; the subsequent light body bands with two distinct indentations on each side, fused in the middle in the last third of the body. Ventral surface of head and neck yellowish cream, belly cream and greyish cream in the last third part of body and on lower tail surface; the dark dorsal bands in part extending towards the venter, forming complete dark bands around the posterior body and on the tail; lateral side of the head cream below, with the lighter pattern beginning in the supralabial region; tip of lower jaw and infralabial region in part greyish brown; dorsal surface of the head and upper head sides a bit paler than the remaining head dorsum. Variation of the new specimens: The Chinese specimen (CIB 115609) largely matches the morphological data of the original description of Lycodon pictus, except for having (1) a slightly shorter tail, TaL/TL 0.210 vs 0.215 in the single female paratype; (2) fewer infralabials 9/10 vs 10; (3) fewer anterior temporals 1 vs 2; however, Janssen et al. (2019) already documented the condition “2*” each for one side of both paratypes, meaning two anterior temporals with the lowermost not touching the postocular, thus referred to as posterior temporal herein, resulting in only one anterior temporal in afore mentioned cases in the type series already. The Vietnamese specimen (VNMN011227) generally agrees with the morphological data of the original description of Lycodon pictus, except for having (1) a slightly longer tail, TaL/TL 0.219 vs 0.211 to 0.215 in the single juvenile and the single female from the type series for which TaL/TL ratio was available, (2) slightly fewer maxillary teeth (12 vs 13 or 14) and (3) fewer subcaudals 85 vs 90 or 91. Both specimens showed more bands on the tail; 14 dark bands in CIB 115609 and 15 dark bands in VNMN011227 (vs 9 and 13); 14 light bands in CIB 115609 and 16 light bands in VNMN011227 (vs 9+ and 13). Extended diagnosis: Lycodon pictus can be differentiated from its congeners by the following morphological characters: dorsal scales in 17–17–15 rows, all smooth; supralabials eight (rarely nine); infralabials ten (rarely nine); one elongated loreal on each side, either in contact with the eye or separated from it; precloacal plate single; ventral scales 212-218 (plus one or two preventral scales); subcaudals 85-91; a total length of 597+ mm in males and 543-591 mm in females; tail / total length ratio 0.210 -0.219 in females; maxillary teeth 12 to 14; dorsal surface of body with 28 or 29 light body bands; dorsal surface of tail with 9 to 15 dark bands, and 13 to 16 cream bands forming a distinct blotch in the vertebral region; ventral surface of body and tail mostly cream with the dark body bands in part extending towards the venter, sometimes forming complete dark bands around the body. Distribution: Lycodon pictus is currently known from the type locality in Cao Bang Province, northern Vietnam and Longzhou County in Guangxi, southern China (Fig. 4). Etymology: The specific name of the species, “ pictus ”, meaning painted or decorated in Latin, refers to the unique dorsal color pattern of this species (Janssen et al., 2019). Because this species is herein reported as a new member of the Chinese snake fauna, we suggest “Jin Bai Huan She (DZ白ẇffi)” as its Chinese common name, deriving from its scientific name. Natural history: Lycodon pictus inhabits karst environment in between elevations of 233 m (Nonggang, Guangxi, China, this study, Fig. 5) and 701 m a.s.l. (type locality; Janssen et al., 2019). In southern China, the species was found at night between 00:00 and 1:00 am after light rain, active in a drainage ditch along a forest path, covered with dead leaves. The surrounding habitat was secondary karst forest, consisting of short hardwood, shrubs and vines. The snake did not attempt to bite when it was handled, neither aggressive nor defensive behavior was observed. In terms of the herpetofauna, several other amphibians and reptiles were observed during the survey period in 2019 in the same microhabitat, including Kurixalus odontotarsus (Ye & Fei in Ye et al., 1993), Rhacophorus kio (Ohler & Delorme, 2006), Draco maculatus (Gray, 1845), Acanthosaura lepidogaster (Cuvier, 1829), Goniurosaurus luii Grismer, Viets & Boyle, 1999, and Psammodynastes pulverulentus (Boie, 1827).

Achalinus tranganensis sp. nov. Figs. 2, 3 Holotype. VNUF R.2018.21 (field number TA.18.21), adult female, from limestone forest (20��15.266���N, 105��53.677���E, 9 m a.s.l.), near Tran Temple, Trang An Landscape Complex, Ninh Binh Province, northern Vietnam, collected by V. Q. Luu, O. V. Lo, N. T. Nguyen on 31 May 2018. Diagnosis. Achalinus tranganensis sp. nov is characterized by the following combination of morphological characters: 1) maxillary teeth 29; 2) suture between the internasals distinctly longer than that between the prefrontals; 3) loreal distinctly wider than high, extending from the nasal to the eye; 4) dorsal scales in 25���23���23 rows, keeled; 5) two pairs of prefrontals; 6) supralabials six; 7) infralabials six; 8) temporals 2+3, the two anterior temporals in contact with eye; 9) ventrals 171; 10) subcaudals 73+, entire; 11) cloacal entire; 12) dorsum in preservative reddish to greyish brown above; 13) the lower part of head side and chin region somewhat paler than remaining head; 14) venter greyish cream, with the anterior region of each ventral and subcaudal somewhat darker. Description of the holotype. Total length 448 mm (SVL 334 mm, TailL 114 mm); tail long, tail tip lost, tail length/ total length ratio 0.25; body slender, cylindrical; head length 9.9 mm; head slightly distinct from neck, dorsally covered with large shields; eye small, with vertically subelliptic pupil; left maxilla with 29 equally sized, curved teeth. Rostral small, triangular, not visible from above; suture between internasals (2.3 mm) longer than that between prefrontals (1.6 mm); nostril in anterior part of nasal; frontal heptagonal, heart-shaped, slightly broader than long, pointed backwards, much shorter than parietals; parietal bordered behind by an elongated nuchal, nuchals separated from each other by three small, elongated scales; one loreal, distinctly longer than high, extending from nasal to eye; medial pair of prefrontals elongated, in broad contact with larger lateral prefrontal; lateral prefrontals in contact with nasal, internasal, medial prefrontal, frontal, supraocular, and loreal; one supraocular; two anterior temporals, both touching eye; two large posterior temporals (a minute scale present between upper anterior and posterior temporals on right); supralabials six, first smallest, fourth and fifth in contact with eye, sixth longest; one curved rectangle mental; six infralabials, first pair in contact with each other, first pair of chin shields larger than posterior, in contact with first three infralabials; posterior pair of chin shields smaller, in contact with third and fourth infralabials. Dorsal scales elliptical, keeled from the neck region posteriorly, in 25-23-23 rows; ventrals 171 (no preventrals), approximately 3 times wider than long, laterally rounded; subcaudals 73+, unpaired; cloacal entire. Colouration in preservative. Reddish to greyish brown above with weakly iridescent colouration in all scales; lower sides of head somewhat paler; venter greyish cream, with anterior region of the ventrals and subcaudals somewhat darker; underside of the tail somewhat darker than belly and the chin region somewhat paler than remaining head (Fig. 4). Colouration in life. Dorsum iridescent reddish to greyish brown; underside of head light brown; venter greyish cream; under tail dark brown. Comparisons. The condition of the prefrontals being arranged in two pairs is unique in Achalinus and thus distinguishes the new species from all other congeners and thus is not repeated in the detailed comparisons in the following. At first glance, Achalinus tranganensis sp. nov. resembles A. juliani in color pattern and dorsal scalerow formula. However, A. tranganensis can be distinguished from A. juliani by having fewer ventrals in the female (171 versus 179); from A. ater by having more anterior temporals (3 versus 2), more dorsal scale rows anteriorly (25 versus 23 or 21), and more subcaudals (73+ versus 47���70); from A. emilyae by having more infralabials (6 ver-sus 5), more posterior temporals (3 versus 2), more dorsal scale rows anteriorly (25 versus 23), more ventrals (171 versus 157���161), and more subcaudals (73+ versus 63); from A. formosanus by having more maxillary teeth (29 versus 14���17), the internasal suture distinctly longer than prefrontal suture (versus internasal suture almost as long as prefrontal suture), the loreal unfused with prefrontal (versus fused), fewer dorsal scale rows (25���23���23 versus 25���29��� 25���27���25), more posterior temporals (3 versus 2), and lacking a black mid-dorsal line; from A. jinggangensis Zong & Ma, 1983 by having more maxillary teeth (29 versus 22), loreal unfused with prefrontal (versus fused), more posterior temporals (3 versus 2), more ventrals (171 versus 156���164), and more subcaudals (73+ versus 51���64); from A. hainanus Huang, 1975 by having more infralabials (6 versus 5), more temporals (2+3 versus 1+2), more dorsal scale rows anteriorly (25 versus 23), more ventrals (171 versus 165���168), and more subcaudals (73+ versus 67���69); from A. meiguensis by having more maxillary teeth (29 versus 17), more dorsal scale rows anteriorly and posteriorly (25 versus 21 or 23; 23 versus 19, respectively), and more subcaudals (73+ versus 39���62); from A. niger by having the internasal suture being distinctly longer than that between the prefrontals (versus almost as long as or shorter), fewer midbody scale rows (23 versus 25), more subcaudals in females (73 versus 52���58), and keeled dorsal scales (versus smooth on anterior part of body); from A. pingbianensis by having the loreal unfused with prefrontal (versus fused), more dorsal scale rows anteriorly (25 versus 23), more ventrals (171 versus 164+2), and more subcaudals (73+ versus 56); from A. rufescens by having more infralabials (6 versus 5), more posterior temporals (3 versus 2), more ventrals in females (171 versus 148���158), and more subcaudals (73+ versus 54���61); from A. spinalis by having more maxillary teeth (29 versus 16���20), the internasal suture being distinctly longer than that between the prefrontals (versus as long as or shorter), more posterior temporals (3 versus 2), more dorsal scale rows anteriorly (25 versus 21���24), and more subcaudals (73 versus 39���67); from Achalinus timi by having the loreal unfused with prefrontal (versus fused), more posterior temporals (3 versus 2), and fewer scale rows at midbody (23 versus 25); from A. werneri by the internasal suture being distinctly longer than that between the prefrontals (versus almost as long as), having more posterior temporals (3 versus 2), and by lacking a black mid-dorsal line; from A. yunkaiensis by having more maxillary teeth (29 versus 20���22), the internasal suture being distinctly longer than that between the prefrontals (versus as long as), more posterior temporals (3 versus 1), more dorsal scale rows anteriorly (25 versus 23), more ventrals (171 versus 151���162), and more subcaudals (73+ versus 38���56). Distribution. Achalinus tranganensis sp. nov. is currently known only from the type locality in the Trang An Landscape Complex, Ninh Binh Province, Vietnam (Fig. 4). Etymology: We name this species after its type locality, the Trang An Landscape Complex, where the new Achalinus was discovered and propose the following common names: Trang An Burrowing Snake (English), R���n xe đi���u tr��ng an (Vietnamese). Natural history. The holotype was found at 2045 h, while crawling on a forest path under dry leaves, at an elevation of 9 m a.s.l. The surrounding habitat is secondary karst forest (Fig. 5). The humidity at the time of collection was approximately 78%, and the air temperature was 29 oC. Other amphibians and reptiles were observed and collected in the area such as Theloderma annae Nguyen, Pham, Nguyen, Ngo, and Ziegler; Sylvirana guentheri (Boulenger), Lycodon futsingensis (Pope), L. subcinctus Boie, Oligodon chinensis (G��nther)., Published as part of Luu, Vinh Quang, Ziegler, Thomas, Ha, Nghia Van, Lo, Oanh Van, Hoang, Tuoi Thi, Ngo, Hanh Thi, Le, Minh Duc, Tran, Dung Hoang & Nguyen, Truong Quang, 2020, A new species of Achalinus (Squamata: Xenodermidae) from Trang An Landscape Complex, Ninh Binh Province, Vietnam, pp. 174-184 in Zootaxa 4877 (1) on pages 177-180, DOI: 10.11646/zootaxa.4877.1.8, http://zenodo.org/record/4572461, {"references":["Zong, Y. & Ma, J. (1983) A new species of the genus Achalinopsis from Jiangxi and the restoration of this genus. Acta Herpetologica Sinica, 2 (2), 61 - 63. [in Chinese]"]}

Redescription of Cyrtodactylus bansocensis Luu, Nguyen, Le, Bonkowski & Ziegler, 2016 including the first record of a female C. bansocensis was described based on two adult males from Khammouane Province (Luu et al. 2016c). Based on the new specimen collected in about 11 km distance from the type locality, including for the first time a female individual, the specimen is described in the following including an expanded diagnosis of the species. Specimen examined. One adult female VNUF R.2016.4 (Fig. 6) collected by Vinh Quang Luu in February 2016 from Khammouane Province, Ban Soi (17��24N, 105��40E, at an elevation of 174 m a.s.l.). Morphological characters. Adult female, SVL 63.8 mm; tail length 94.6 mm. The examined specimen differs from the original description of Luu et al. (2016c) by having more supralabials and infralabials (11 or 12 vs 8���10; 10 vs. 8, respectively), more dorsal tubercle rows (20 vs. 14 or 15) and more scales around midbody (102 vs. 86 or 87). For further morphological data see Table 6. Expanded diagnosis. As the original description of Cyrtodactylus bansocensis was based on two adult males (Luu et al. 2016c) we herein expand the diagnosis of this species as follows: medium size, SVL reaching 74.0 mm; dorsal pattern with four light transverse bands between limb insertions; supranasals in contact; 14���20 irregular dorsal tubercle rows at midbody; lateral skin fold present without tubercles; 34 or 35 ventral scale rows between ventrolateral folds; 34 precloacal and femoral pores in a continuous row in males (or interrupted by poreless scales); 16 precloacal and femoral pores in a continuous row in females (two outer femoral pores just pitted on each side), enlarged femoral and precloacal scales present; 4���7 postcloacal tubercles on each side; dorsal tubercles present at base of tail; subcaudal scales transversely enlarged (modified after Luu et al. 2016c). Natural history notes. The female specimen was collected at night (20:20) on a limestone cliff, ca. 1.5 m from the ground. The surrounding habitat was karst forest. The relative humidity was 52% and the air temperature was 25.5��C. Distribution. The female specimen of Cyrtodactylus bansocensis was found close to the type locality in Khammouane Province, Laos., Published as part of Schneider, Nicole, Luu, Vinh Quang, Sitthivong, Saly, Teyni��, Alexandre, Le, Minh Duc, Nguyen, Truong Quang & Ziegler, Thomas, 2020, Two new species of Cyrtodactylus (Squamata: Gekkonidae) from northern Laos including new finding and expanded diagnosis of C. bansocensis, pp. 503-530 in Zootaxa 4822 (4) on pages 520-521, DOI: 10.11646/zootaxa.4822.4.3, http://zenodo.org/record/4452362, {"references":["Luu, V. Q., Bonkowski, M., Nguyen, T. Q., Le, M. D., Schneider, N., Ngo, H. T. & Ziegler, T. (2016 a) Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos. Zootaxa, 4107 (2), 101 - 140. https: // doi. org / 10.11646 / zootaxa. 4162.2.4","Luu, V. Q., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016 c) A new species of karst-dwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4079 (1), 87 - 102. https: // doi. org / 10.11646 / zootaxa. 4079.1.6"]}

Cyrtodactylus houaphanensis sp. nov. (Fig. 2) Holotype. One adult male, IEBR A.2013.109, collected by Alexandre Teynié in June 2013 from Houaphan Province, near Viengxai, at an elevation of 850 m a.s.l. Diagnosis. The new species can be distinguished from all its congeners by a unique combination of characters: SVL 75.8 mm; supralabials 9 or 10; infralabials 8 or 9; ventral scales 35; dorsal tubercles in 20 rows at midbody; precloacal pores in the male 6; femoral pores absent; subcaudals enlarged; five irregular, brown bands between limb insertions. Description of holotype. Adult male, SVL 75.8 mm; tail regenerated (TaL 59.1 mm); head depressed (HL/HW 1.4), distinct from neck; snout scales small, homogeneous, granular, larger than those in frontal and parietal regions; rostral wider than high (RW 3.8 mm, RH 2.6 mm), square-shaped, medially with a straight, vertical suture, in contact with nasorostral, nare, and first supralabial on each side; internasal single; nares oval, surrounded by supranasal, rostral, first supralabial, and three postnasals; supralabials 9 or 10; ears oval; mental triangular, slightly narrower than rostral (MW 3.3 mm), in contact with two enlarged postmentals and the first infralabial on each side, postmentals surrounded by first infralabial on each side (and second on right side) and seven granular scales posteriorly, two outer ones enlarged; infralabials 8 or 9; gular scales granular; scales between fifth supralabials across the dorsal head surface in 43 rows; scales between anterior corners of eye orbits 31; interorbital region with small, round or oval, convex scales; dorsal surface of head with enlarged tubercles; pupil vertical (Fig. 3 a–d). Dorsal scales granular to flattened; dorsal tubercles on head, back and tail base conical, each surrounded by 8 or 9 granular scales, in 20 irregular longitudinal rows at midbody; scales around midbody 127; ventral scales smooth, medial scales 2 or 3 times larger than dorsal scales, round or oval, in 35 longitudinal rows at midbody; scales between mental and cloacal slit 184; ventrolateral folds present; precloacal groove absent; femoral scales enlarged, femoral pores absent; precloacal scales enlarged, precloacal pores 6; postcloacal tubercles 2 on each side; subcaudals enlarged (Fig. 3e); tubercles on forelimbs absent, present on hindlimbs; fingers and toes free of webbing; lamellae under fourth finger 17, under fourth toe 19–21 (Table 1). Coloration in life: Ground color of dorsal head and back dark brown with a yellow pattern; a discontinuous nuchal loop, extending from posterior corner of the eye above tympanum to the neck, dark brown, ending in two dark spots; head with dark brown blotches; dorsum with five irregular and sometimes interrupted dark brown bands on the back between limb insertions; ventral surface yellowish-beige; dorsal surface of fore and hind limbs with irregular dark bands; regenerated tail with irregular dark brown and beige pattern (Fig. 2). Comparisons. We compared the new species with its congeners from Laos (Table 2) and with other Cyrtodactylus from the Indochina region including Vietnam, Thailand and Cambodia (Table 3). Morphologically, C. houaphanensis sp. nov. resembles the members of the C. wayakonei group due to the low amount of precloacal pores. Therefore, below we compared the new species with members of the C. wayakonei species group. Morphologically, C. houaphanensis sp. nov. is most similar to the genetically only distantly related C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau from Vietnam but can be distinguished from the latter species in morphology by its smaller size (SVL 75.8 mm vs. 91–99.3 mm), having generally fewer infralabials (8 or 9 vs. 9–11), and the presence of an internasal scale which is absent in C. chauquangensis (Hoang et al. 2007). From its sister species C. puhuensis Nguyen, Yang, Le, Nguyen, Orlov, Hoang, Nguyen, Jin, Rao, Hoang, Che, Murphy & Zhang, the new species differs by the absence of enlarged femoral scales (vs. presence), and having slightly more precloacal pores in males (6 vs. 5). The new species has enlarged subcaudal scales and thus differs from the following species which are lacking enlarged subcaudals: C. bobrovi Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler; C. otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler; C. vilaphongi Schneider, Nguyen, Le, Nophaseud, Bonkowski & Ziegler; C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler. The new species is lacking femoral pores in males and thus differs from the following species which have femoral pores in males: C. huongsonensis Luu, Nguyen, Do & Ziegler; C. soni Le, Nguyen, Le & Ziegler; C. sonlaensis Nguyen, Pham, Ziegler, Ngo & Le; C. muangfuangensis Sitthivong, Luu, Ha, Nguyen, Le & Ziegler; and the new Cyrtodactylus species described below. ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page The new species differs from C. cucphuongensis Ngo & Chan by the absence of enlarged femoral scales (vs. present), the presence of precloacal pores in males (vs. absent), and having fewer ventral scales (35 vs. 42); from C. martini Ngo by the absence of enlarged femoral scales (vs. present), having more precloacal pores in males (6 vs. 4), and having fewer ventral scales (35 vs. 39–43); from C. spelaeus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov by having fewer precloacal pores in males (6 vs. 8–9), by its smaller size (75.8 mm vs. 91 mm) and by having a different color pattern of dorsum (banded vs. blotched); from C. taybacensis Pham, Le, Ngo, Ziegler & Nguyen by its smaller size (maximum SVL 75.8 mm vs. 97.5), the absence of enlarged femoral scales (vs. present), and having fewer precloacal pores in males (6 vs. 11–13). Natural history notes. The holotype was found on a large rock partially covered with vegetation 1.2 m above the ground. A second specimen was observed active at the bottom of a rock cavity 2 m above the ground. Both specimens were discovered on a steep rock at night (20:30–20:45) during the rainy season at the foot of a limestone cliff of the karst formations. The adjacent lowland was mainly covered with various small-scale agricultural crops, patches of secondary forests and some scrub and grasslands, where no large primary forest remained. Distribution. Cyrtodactylus houaphanensis sp. nov. is currently known only from the type locality in Houaphan Province, Laos. Etymology. We name this species after its type locality in Houaphan Province, Laos and propose Houaphan Bent-toed Gecko as a common name.

Cyrtodactylus ngoiensis sp. nov. (Fig. 4) Holotype. One adult male, IEBR 4548 (Fig 4a), collected by Alexandre Teyni�� in 2012 from Luang Prabang Province, Ngoi District (at an elevation of 420 m a.s.l.). Paratypes. Three adult females: ZFMK 102768, IEBR A.2013.110 (Fig. 4b), IEBR A.2013.111, collected by Truong Quang Nguyen and Alexandre Teyni�� in August 2013 from Luang Prabang Province, Ngoi District (at elevations of 402���427 m a.s.l.). Diagnosis. The new species can be distinguished from all its congeners on the basis of the following combination of characters: maximum SVL 95.3 mm; supralabials 6���9; infralabials 8���11; ventral scales 38���43; dorsal tubercles in 15���21 rows at midbody; enlarged femoral scales present; precloacal pores 7 in the male, 7 pitted scales in females; femoral pores 14 in the male, absent in females; five bright yellow transverse dorsal bands; subcaudals enlarged. Description of holotype. Adult male, SVL 88.8 mm, tail broken; head depressed (HL/HW 1.4), distinct from neck; snout scales small, homogeneous, granular, larger than those in frontal and parietal regions; rostral wider than high, square-shaped, medially with a straight, vertical suture, in contact with nasorostral, nare, and first supralabial on each side; internasal absent; nares round or oval, surrounded by supranasal, rostral, first supralabial, and two or three enlarged postnasals; supralabials 8 or 9; ear in oval shape; mental triangular, slightly narrower than rostral, in contact with two postmentals and the first infralabial on each side, postmentals surrounded by first infralabial on each side and 10 granular scales posteriorly, two outer ones enlarged; infralabials 8 or 9; gular scales granular; scales between fifth supralabials across the dorsal head surface in 50 rows; scales between anterior corners of eye orbits 32; interorbital region with small, round or oval, convex scales; dorsal postorbital surface of head with enlarged tubercles; pupil vertical (Fig. 5 a���d). Dorsal scales granular to flattened; dorsal tubercles conical, present on head, back and tail base, each surrounded by 8���10 granular scales, in 21 irregular longitudinal rows at midbody; scales around midbody 102; ventral scales smooth, medial scales 2 or 3 times larger than dorsal scales, round, in 38 longitudinal rows at midbody; scales between mental and cloacal slit 179; ventrolateral folds slightly developed; precloacal groove absent; femoral scales enlarged, pitted femoral scales 14; precloacal scales enlarged, precloacal pores 7; postcloacal tubercles 3 on each side (Fig. 5e); tubercles absent on forelimbs, present on hindlimbs; fingers and toes free of webbing; lamellae under fourth finger 16 or 17, under fourth toe 20 (Tables 4 and 5). Coloration in life: Ground coloration of dorsal head and back brown; head with dark brown spots; nuchal loop U-shaped, sometimes broken, from posterior corner of eye over the tympanum to the neck, dark brown, edged in bright yellow; five brown transverse bands between limb insertions, edged in bright yellow, darker in center; dorsal surface of fore and hind limbs with alternating brown and dark brown pattern; tail brown dorsally with nine to ten light brown bands; ventral surface beige to cream. Variation. Sexual dimorphism is shown in adult females lacking femoral pores (vs. 14 pitted scales in the male); in addition, females had fewer dorsal tubercle rows (15���19) and more scales from mental to cloacal slit (192���199); a single internasal present in the female ZFMK 102768; scales between fifth supralabials across the head 39���64. Female specimens show enlarged subcaudal scales, which is not determinable in male holotype due to missing tail. Comparisons. We compared the new species with its congeners from Laos (Table 2) and with other Cyrtodactylus from the Indochina region including Vietnam, Thailand and Cambodia (Table 3). Morphologically, Cyrtodactylus ngoiensis sp. nov. resembles the members of the C. wayakonei group due to the low amount of precloacal pores (Cyrtodactylus ngoiensis sp. nov.: PP in the male and (or pitted scales) in females: 7). Therefore, below we compared the new species with members of the C. wayakonei species group. The new species has enlarged subcaudal scales and thus differs from the following species which are lacking enlarged subcaudals: C. bobrovi; C. otai; C. vilaphongi; C. wayakonei. The new species has femoral pores in males and thus differs from the following species which are lacking femoral pores in males: C. chauquangensis; C. cucphuongensis; Cyrtodactylus houaphanensis sp. nov.; C. martini; C. puhuensis; C. spelaeus; C. taybacensis. The new species has a contiguous series of precloacal-femoral pores in males and thus differs from the following species, which are separated by poreless scales: C. huongsonensis; C. sonlaensis; C. muangfuangensis. The new species differs from C. soni by having more femoral pores in males (14 vs. 5���8), and the presence of 7 precloacal pitted scales in females (vs. 7���8 pores). Morphologically, Cyrtodactylus ngoiensis sp. nov. is most similar to C. dumnuii from Thailand but the new species can be distinguished from the latter by its larger size (maximum SVL 95.3 vs. 84.2), having more precloacal pores in males (7 vs. 5���6) and pitted precloacal scales in females (7 pitted scales vs. 0���7 pores) (Bauer et al. 2010). In addition, Cyrtodactylus ngoiensis sp. nov. has less pronounced dorsal tubercles than C. dumnuii and a different colour pattern (uniform brown bands in C. dumnuii versus thinner brown transverse bands edged in whitish yellow in Cyrtodactylus ngoiensis sp. nov.). Natural history notes. The female paratype ZFMK 102768 was found on a limestone wall near a cave (at elevation of 402 m a.s.l.), about 1 m above the forest ground. IEBR A.2013.110 and IEBR A.2013.111 were found in a small hole of a limestone wall, about 0.2���0.5 m above the forest ground. Distribution. Cyrtodactylus ngoiensis sp. nov. is currently known only from the type locality in Luang Prabang Province, Laos. Etymology. We name this species after its type locality in Ngoi District, Luang Prabang Province, Laos and propose the common name Ngoi Bent-toed Gecko (English)., Published as part of Schneider, Nicole, Luu, Vinh Quang, Sitthivong, Saly, Teyni��, Alexandre, Le, Minh Duc, Nguyen, Truong Quang & Ziegler, Thomas, 2020, Two new species of Cyrtodactylus (Squamata: Gekkonidae) from northern Laos including new finding and expanded diagnosis of C. bansocensis, pp. 503-530 in Zootaxa 4822 (4) on pages 516-520, DOI: 10.11646/zootaxa.4822.4.3, http://zenodo.org/record/4452362, {"references":["Bauer, A., Kunya, K., Sumontha, M., Niyomwan, P., Pauwels, O. S. G., Chanhome, L. & Kunya, T. (2010) Cyrtodactylus dumnuii (Squamata: Gekkonidae), a new cave-dwelling gecko from Chiang Mai Province, Thailand. Zootaxa, 2570 (1), 41 - 50. https: // doi. org / 10.11646 / zootaxa. 2570.1.2"]}

Key to Laotian species of Cyrtodactylus (referring to Table 2) 1 Precloacal-femoral pores in males in separated series.......................................... C. muangfuangensis - Precloacal-femoral pores in males in contiguous series or precloacal-femoral pores absent in males.................... 2 2 Enlarged subcaudals present............................................................................. 3 - Enlarged subcaudals absent............................................................................ 15 3 Precloacal-femoral pores present in males.................................................................. 4 - Precloacal-femoral pores absent or unknown in males....................................................... 19 4 Dorsum blotched..................................................................................... 5 - Dorsum banded....................................................................................... 6 5 Precloacal pores absent in females, Supralabials 11���16................................................ C. jarujini - Precloacal pores in females 0���14, Supralabials 9���11................................................ C. multiporus 6 Ventrals 27���29................................................................................ C. rufford - Ventrals 31 or more................................................................................... 7 7 Precloacal-femoral pores in males 20���29................................................................... 8 - Precloacal-femoral pores in males 34 or more............................................................... 9 8 Precloacal pores (pitted scales) in females present................................................ C. sommerladi - Precloacal pores in females absent............................................................. C. soudthichaki 9 Precloacal-femoral pores in males 34, Midbody scales 86���87....................................... C. bansocensis - Precloacal-femoral pores in males 35 or more.............................................................. 10 10 Precloacal-femoral pores in females 0���34................................................................. 11 - Precloacal-femoral pores in females 38............................................................ C. calamei 11 Midbody scales 155���172.............................................................................. 12 - Midbody scales 179 or more........................................................................... 13 12 Ventrals 31���32, Precloacal-femoral pores in males 44, in females 21...................................... C. jaegeri - Ventrals 32���38, Precloacal-femoral pores in males 40��� 44, in females 0���17........................ C. khammouanensis 13 SVL ��� 71.2mm, Dorsal tubercle rows 19���24..................................................... C. lomyenensis - SVL> 72mm, Dorsal tubercle rows 14���20................................................................ 14 14 Dorsal tubercle rows 16���20, Precloacal-femoral pores in females 24���34.................................. C. darevskii - Dorsal tubercle rows 14���19, Precloacal-femoral pores in females 0���28.............................. C. hinnamnoensis 15 Precloacal pores in females present...................................................................... 16 - Precloacal pores in females absent....................................................................... 17 16 Dorsum banded............................................................................. C. wayakonei - Dorsum blotched.................................................................. C. pseudoquadrivirgatus 17 Dorsum banded............................................................................. C. vilaphongi - Dorsum blotched.................................................................................... 18 18 Ventrals 30.................................................................................. C. buchardi - Ventrals 40���50................................................................................ C. cryptus 19 Dorsum blotched.................................................................................... 20 - Dorsum banded...................................................................................... 21 20 Precloacal pores present in females................................................................ C. teyniei - Precloacal pores absent in females................................................................ C. spelaeus 21 Femoral pores present in males......................................................................... 22 - Femoral pores absent in males.......................................................................... 24 22 Precloacal pores in males 7���10......................................................................... 23 - Precloacal pores in males 14................................................................. C. interdigitalis 23 Precloacal pores in females 7............................................................ C. ngoiensis sp. nov. - Precloacal pores in females 10-12............................................................ C. thathomensis 24 Precloacal pores in males 4, in females 4���6........................................................... C. pageli - Precloacal pores in males 6......................................................... C. houaphanensis sp. nov., Published as part of Schneider, Nicole, Luu, Vinh Quang, Sitthivong, Saly, Teyni��, Alexandre, Le, Minh Duc, Nguyen, Truong Quang & Ziegler, Thomas, 2020, Two new species of Cyrtodactylus (Squamata: Gekkonidae) from northern Laos including new finding and expanded diagnosis of C. bansocensis, pp. 503-530 in Zootaxa 4822 (4) on pages 521-522, DOI: 10.11646/zootaxa.4822.4.3, http://zenodo.org/record/4452362

Two new Cyrtodactylus species are described from Houaphan and Luang Prabang provinces in Laos based on morphological and molecular data. Cyrtodactylus houaphanensis sp. nov. differs from all other Cyrtodactylus in the C. wayakonei species group by at least 3.3 % genetic divergence in the COI gene and can be diagnosed in morphology as follows: SVL 75.8 mm; supralabials 9 or 10; infralabials 8 or 9; ventral scales 35; dorsal tubercles in 20 rows at midbody; precloacal pores 6 in the male; femoral pores absent; subcaudals enlarged; five irregular, brown bands between limb insertions. The new species morphologically resembles C. chauquangensis and revealed to be a sister taxon to C. puhuensis according to our genetic analyses, from which it mainly differs in the absence of enlarged femoral scales. Cyrtodactylus ngoiensis sp. nov. differs from other closely related congeners by at least 11.6 % genetic divergence in the COI gene and can be diagnosed in morphology as follows: maximum SVL 95.3 mm; supralabials 6–9; infralabials 8–11; ventral scales 38–43; dorsal tubercles in 15–21 rows at midbody; enlarged femoral scales present; precloacal pores 7 in the male, 7 pitted scales in females; femoral pores 14 in the male, absent in females; five bright yellowish transverse dorsal bands; subcaudals enlarged. The new species is shown to be a member of the C. wayakonei species group, but morphologically resembles C. dumnuii from Thailand. According to our phylogenetic analyses, it is a basal taxon to a clade comprising C. spelaeus, C. chauquangensis, C. vilaphongi, C. cucphuongensis, C. puhuensis, C. houaphanensis, C. otai and C. bobrovi. Based on a new record of C. bansocensis from central Laos, which represents the first recorded female we provide a redescription of C. bansocensis including expanded diagnosis. In addition, an updated identification key for the Cyrtodactylus known from Laos is provided.

Hemiphyllodactylus nahangensis sp. nov. (Figs. 3, 4) Holotype. IEBR 4741 (Field number TQ.2018.140), adult male, collected on 20 June 2018 by C. T. Pham, A.M. Luong, T. Q. Phan and H. T. Ninh from the limestone karst area near Trung Phin Village (20 o 30.215’N, 105 o 23.402’E, at an elevation of 915 m a.s.l.), Sinh Long Commune, Na Hang District, Tuyen Quang Province, northern Vietnam. Paratypes. Two specimens collected from the same site as the holotype: IEBR 4742 (Field number TQ.2018.141), adult male, collected on 20 June 2018 and IEBR 4743 (Field number TQ.2018.96), adult female, collected on 18 June 2018, by C. T. Pham, A.M. Luong, T. Q. Phan and H. T. Ninh. Diagnosis. The new species from northern Vietnam differs from the remaining congeners of the genus Hemiphyllodactylus by a combination of the following characters: a bisexual taxon; SVL of adults 41.4–43.6 mm; dorsal scale rows 18–23; ventral scale rows 9–13; chin scales bordering mental and first infralabial, distinctly enlarged; digital lamellae formula 3-4-5-4 (forefoot) and 4-5-5-5 (hindfoot); 22–24 pore-bearing femoral and precloacal scales, in a continuous row, absent in females; cloacal spur single in both sexes; dark lateral head stripe indistinct; postsacral mark cream and bearing anteriorly projecting arms. Description of holotype. Adult male; size small (SVL 43.6 mm); tail length (TaL 37.4 mm); body elongate (TrunkL 21.3 mm); head longer than wide (HeadL 10.7 mm, HeadW 7.4 mm), not markedly depressed (HeadD/ HeadL 0.34), distinct from neck; eye large, pupils vertical (EyeD 2.5 mm); ear oval shaped, small (EarD 0.69 mm); nare–eye length (NarEye 3.1 mm), snout-eye length (SnEye 3.8 mm), internarial distance (SnW 1.3 mm). Proportions: Trunk/SVL 0.44, HeadL/SVL 0.24, HeadW/SVL 0.17, HeadW/HeadL 0.69, HeadD/HeadL 0.34, SnEye/HeadL 0.36, NarEye/HeadL 0.29, EyeD/HeadL 0.24, EyeD/SnEye 0.67, EyeEar/EyeD 1.39, SnW/HeadL 0.12, EyeD/NarEye 0.82, SnW/HeadW 0.18. Scalation: Rostral very large, wider than high, with a shallow suture, bordered posteriorly by large supranasals; supralabials 11/11, gradually decreased in size towards angle of jaw; nare in contact with rostral anteriorly, first supralabial ventrally, supranasal dorsally and three nasals posteriorly on each side; supranasals separated from each other by four small granular internasals; snout flat; pupil vertical, upper eyelid with small supraciliaries; ear oval-shaped, approximately 27% of the eye diameter, not bordered by enlarged scales; infralabials 10/9; mental triangular, slightly wider than rostral, bordered by the first infralabial on each side and posteriorly by two enlarged postmentals; eight chin scales touching internal edge of infralabials and mental between the juncture of the second and third infralabials on each side of the head, anterior pair enlarged; gular scales small, subimbricate; throat and pectoral scales which grade into slightly larger, subimbricate; dorsal scales small, granular, in 22 rows at midbody (contained within one eye diameter); ventrolateral folds absent; enlarged turbercles absent; ventral scales flat, subcircular, larger than dorsal scales, in 9 rows at midbody on venter (contained within one eye diameter); femoral pores and precloacal pores in continuous series, 24 in total; cloacal spur single; dorsal surface of fore and hindlimbs covered with granular scales; terminal two phalanges free, claws absent on first finger and on first toe, present on second to fifth digit of fore and hindfoot; pads of digits II–V each with large triangular lamella, digital formula 3-4-5-4 (forefoot) and 4-5-5-5 (hindfoot); lamellae three on first fingers, three on first toes; caudal scales flat, not forming distinct caudal segments. Coloration in life: Ground color of dorsum, head and limbs grey with indistinct black markings; dark lateral head stripe indistinct; light postorbital stripe extending to anterior margin of forearm; ventral surface cream with dark mottling; dorsum with indistinct dark pattern, upper zone of flank with light spots and indistinct dark longitudinal markings; postsacral mark cream in U-shape and bearing anteriorly projecting arms; original tail with some dark spots, edged by light grey and cream markings. Coloration in preservative: Color became darker, dark pattern disappeared on dorsum, head and dorsal surface of limbs. Tail pattern did not change noticeably in preservation. Sexual dimorphism and variation. Measurements and scalation characters of the paratypes are given in Table 2. The female differs from the males in the absence of hemipenial swellings at the tail base. The scale counts vary among the type series: scales between supranasals 4–6; supralabials 10–12; infralabials 9–11; chin scales 8 or 9; dorsal scale rows 18–23; ventral scale rows 9–13. The males have 22–24 femoral and precloacal pores (versus absent in female). Ground color on upper surface of head, body, and tail is also different among individuals, varying from whitish grey to brownish grey with yellowish marks. Distribution. Hemiphyllodactylus nahangensis sp. nov. is currently known only from the type locality in Na Hang District, Tuyen Quang Province, northern Vietnam (Fig.1). Etymology. Specific epithet nahangensis is a toponym in reference to the type locality of the species. For the common names we suggest Nahang Slender Gecko (English) and Thạch sùng dẹp na hang (Vietnamese). Natural history. The new species is associated with residential habitat. Type specimens were collected between 20:00 and 22:00 h, one individual was collected on the wooden wall of a house in Trung Phin Village, two individuals were collected on a branch of a peach tree near the house, approximately 2.5–3.5 m above the ground. The surrounding habitat of Trung Phin village was disturbed evergreen karst forest of medium, small hardwood and shrub trees. Air temperature ranged from 25ºC to 31ºC and relative humidity was 63–87%. Comparisons: We compared the new species from northern Vietnam with all other members of the genus Hemiphyllodactylus from Vietnam, Laos, and China based on examination of specimens (see Appendix) and data obtained from the literature (Boulenger 1903; Barbour 1924; Smith 1935; Taylor 1963; Zhou et al. 1981 with English translation of Ota 1996; Bourret 2009; Zug 2010; Grismer et al. 2013, 2014a,b, 2015, 2018a,b, 2020; Nguyen et al. 2013, 2014, 2020; Ngo et al. 2014; Guo et al. 2015; Cobos et al. 2016; Yan et al. 2016; Sukprasert et al. 2018; Sung et al. 2018; Eliades et al. 2019; Agarwal et al. 2019, 2020). For comparisons with other species of Hemiphyllodactylus see Table 3. Hemiphyllodactylus nahangensis sp. nov. differs from H. banaensis Ngo, Grismer, Pham & Wood by having a smaller size (maximum SVL 43.6 mm versus 51 mm in H. banaensis), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 5 and 5, respectively, in H. banaensis), and more precloacal and femoral pores in males (22–24 versus 18–21 in H. banaensis); from H. bonkowskii by having more chin scales (8–9 versus 5–7 in H. bonkowskii), fewer dorsal scale rows (18–23 versus 24–27 in H. bonkowskii), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 5 and 4–5, respectively, in H. bonkowskii), digital lamellae formula 3-4-5-4 (forefoot) and 4-5-5- 5 (hindfoot) (versus 3-4-4-4 and 4-5-5-4, respectively, in H. bonkowskii), and more precloacal and femoral pores in males (22–24 versus 19 in H. bonkowskii); from H. dushanensis by having a smaller size (maximum SVL 43.6 mm versus 50.6 mm in H. dushanensis), more dorsal scale rows (18–23 versus 11–15 in H. dushanensis), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 4–5 and 5–6, respectively, in H. dushanensis), and the presence of anteriorly projecting arms on postsacral (versus absent in H. dushanensis); from H. hongkongensis by having more chin scales (8–9 versus 5–6 in H. hongkongensis), more dorsal scale rows (18–23 versus 12–15 in H. hongkongensis), fewer subdigital lamellae 3 (1ToeLm) (versus 5 in H. hongkongensis), digital lamellae formula 3-4-5-4 (forefoot) and 4-5-5-5 (hindfoot) (versus 3-4-4-4/3-3-4-4 and 3-4-4-4/4-4-5-4/4-4-5-5, respectively, in H. hongkongensis); from H. huishuiensis Yan, Lin, Guo, Li & Zhou by having more scales between supranasals (4–6 versus 2–3 in H. huishuiensis), more dorsal scale rows (18–23 versus 13–15 in H. huishuiensis), more ventral scale rows (9–13 versus 7–9 in H. huishuiensis), and more precloacal and femoral pores in males (22–24 versus 18–20 in H. huishuiensis); from H. indosobrinus Eliades, Phimmachak, Sivongxay, Siler & Stuart by having more scales between supranasals (4–6 versus 3 in H. indosobrinus), fewer supralabials (10–12 versus 15 in H. indosobrinus), fewer dorsal scale rows (18–23 versus 30 in H. indosobrinus), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 4 and 5, respectively, in H. indosobrinus), and more precloacal and femoral pores in males (22–24 versus 18 in H. indosobrinus); from H. kiziriani by having more scales between supranasals (4–6 versus 2–3 in H. kiziriani), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 5 and 5, respectively, in H. kiziriani), digital lamellae formula 3454 (forefoot) and 4-5-5-5 (hindfoot) (versus 3-4-4-4 and 4-4(5)-4(5)-4, respectively, in H. kiziriani), and more precloacal and femoral pores in males (22–24 versus 10–13 in H. kiziriani); from H. ngocsonensis by having more scales between supranasals (4–6 versus 3 in H. ngocsonensis), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 4–5 and 5–6, respectively, in H. ngocsonensis), digital lamellae formula 3454 (forefoot) and 4555 (hindfoot) (versus 3-4-4-4 and 4-5-5-4, respectively, in H. ngocsonensis), and more precloacal and femoral pores in males (22–24 versus 20 in H. ngocsonensis); from H. serpispecus by having more scales between supranasals (4–6 versus 2 in H. serpispecus), fewer dorsal scale rows (18–23 versus 26 in H. serpispecus), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 4 and 4, respectively, in H. serpispecus), digital lamellae formula 3-4-5-4 (forefoot) and 4-5-5-5 (hindfoot) (versus 3-4-4-4 and 3-4-4-5, respectively, in H. serpispecus), and more precloacal and femoral pores in males (22–24 versus 11 in H. serpispecus); from H. zugi Nguyen, Lehmann, Le, Duong, Bonkowski & Ziegler by having fewer ventral scales (9–13 versus 14–16 in H. zugi), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 4–5 and 4–5, respectively, in H. zugi), digital lamellae formula 3-4-5-4 (forefoot) (versus 3-4-4- 4 in H. zugi), and more precloacal and femoral pores in males (22–24 versus 18–21 in H. zugi).

Dixonius lao sp. nov. (Figs. 2���5) Holotype. Adult male, VNUF R.2016.2 (Field no. TK16.2) in Na Ngua village, Thakhek Town, Khammouane Province (17 o 32N, 104 o 51E, at an elevation of 145 m a.s.l.), collected by V. Q. Luu and N. Schneider on 24 February 2016. Paratypes. IEBR A.2019.5 (Field no. TK16.3), adult female; IEBR A.2019.6 (Field no. TK16.1), subadult female; the same data as the holotype. Diagnosis. A species of the genus Dixonius that can be distinguished from its congeners by a combination of the following characters: maximum SVL 55.4 mm; 20���23 longitudinal rows of dorsal tubercles at midbody; 23 or 24 longitudinal rows of ventrals across the abdomen; 8���10 supralabials, 7 or 8 in mid-orbital position; 7 or 8 infralabials; 8 precloacal pores in male; the male without femoral pores; precloacal and femoral pores absent in females; uniformly pebble brown dorsum. Description of the holotype. Adult male, SVL 50.1 mm with dorsolaterally flattened body, TaL 69.0 mm, trunk length 20.6 mm. Head longer than wide, depressed, distinct from neck. Head length 14.1 mm, head width 9.2 mm, eye of moderate size 3.6 mm, ear-opening oblique (EL 1.4 mm), snout-eye length 5.6 mm, internarial distance 1.7 mm. Rostral very large, wider than high (width 2.2 mm, height 1.2 mm), with distinct suture; supralabials 10/9 (8/ 7 in midorbital position); nostril in contact with rostral, first supralabial, supranasal, and two nasals posteriorly on each side; supranasals in broad contact; without intersupranasals; snout flat, covered with granular scales; pupil vertical; ear opening oblique, oval, approximately one third of the eye diameter, without bordering enlarged scales; infralabials 8/8, decreasing gradually in size; mental triangular, wider than high (width 2.5 mm, height 1.7 mm); two pairs of enlarged postmentals, first pair very large and in contact with each other, second pair about one third to one half size of first, in contact with the first and second infralabials and separated from each other by four gular scales; dorsal tubercle rows at midbody 22, keeled and separated from each other by one smaller scale which is keeled or at least almost conical shaped; ventral scales larger than dorsal scales, in 23 rows at midbody; dorsal surface of fore- and hind limbs covered with shallowly keeled scales, area around limb insertions covered with small granular scales; paravertebral scales, number of scales in a paravertebral row from first scale posterior to parietal scale to last scale at the level of vent opening 43; paravertebral scales in a row between limb insertions 25; lamellae on fourth toe 15; precloacal pores 8, in an angular series; tail covered with smooth scales, in different sizes; subcaudals 42, enlarged and undivided. Coloration in life. In life, dorsum of the holotype pebble brown (Fig. 2); head with brownish olive spots; canthal stripe absent; ventral surface light beige to uniformly whitish as the belly and the throat; supralabials grey with dark grey spots; upper surface of fingers and toes uniformly light grey; dorsal surface of the tail light brownish grey, with uniformly distributed light spots along the tail tip. Sexual dimorphism. Morphologically, the female paratypes corresponded well with the holotype, except for the lack of precloacal pores. Comparisons. We compared the new species from Laos with all other known Dixonius species (see Table 4 for a comparison with most similar species). Morphologically, the new species is most similar to Dixonius minhlei from Vietnam in the number of precloacal pores and ventral scale rows. However, the new species can be distinguished from the latter by having more dorsal tubercle rows at midbody (20���23 versus 14 or 15), the absence of canthal stripe (versus canthal stripe continuing behind orbit to back of head), and by a different color pattern (pebble brown dorsum versus olive grey ground color on dorsum, with more or less discernible brownish olive blotches in D. minhlei); from D. siamensis, which is known from Laos, Cambodia, Thailand and Vietnam, by having more dorsal tubercle rows (20���23 versus 10���14), more precloacal pores (8 versus 6 or 7), and generally more supralabials (8���10 versus 7 or 8); from D. hangseesom from Thailand by having more dorsal tubercle rows (20���23 versus 12���14), more paravertebral scales in a row from first scale posterior to parietal scale to last scale at the level of vent opening (PV 40���43 versus 26), and coloration of tail almost the same as the dorsum (versus orange tail); from D. kaweesaki from Thailand by its larger size (maximum SVL 55.4 mm versus 41.6 mm), having more dorsal tubercle rows at midbody (20���23 versus 12 or 13), and in dorsal head and body pattern (back pebble brown and head with brownish olive spots versus dorsal surface of head gray in D. kaweesaki); from D. melanostictus from Thailand and Vietnam by having more dorsal tubercle rows (20���23 versus 10 or 11), more paravertebral scales in a row between limb insertions (PV��� 24���25 versus 10 or 11), and the absence of canthal stripe (versus canthal stripe extending along flanks in D. melanostictus); from D. aaronbaueri from Vietnam by having more ventral scale rows at mid-body (23 or 24 versus 18 or 19), more dorsal tubercle rows at midbody (20���23 versus 11), more precloacal pores in males (8 versus 5), and different dorsal pattern (back pebble brown versus dorsal yellowish-orange to bright orange color from head to tail tip); from D. taoi from Vietnam by its larger size (maximum SVL 55.4 mm versus 43.9 mm), having more dorsal tubercle rows (20���23 versus 11 or 12), more precloacal pores (8 versus 5 or 6) and different dorsal pattern (back pebble brown versus one or two irregular rows of yellowish marks running from head along flanks in D. taoi); from D. vietnamensis from Vietnam and Cambodia in having more dorsal tubercle rows at mid-body (20���23 versus 16), more supralabials (8���10 versus 5 or 6), more ventral scale rows at midbody (23 or 24 versus 20), more lamellae under fourth toe (15 versus 13), and more precloacal pores (8 versus 5���7). Natural history notes. The type series was found between 20:00 and 21:00h on the forest floor or on karst cliffs, about 0.3 m above the ground. The surrounding habitat was secondary karst forest on the mountain (Fig. 6). The relative humidity ranged from 58% to 70%, and air temperature was 25���26 oC. Distribution. Dixonius lao sp. nov. is currently known only from the type locality in Khammouane Province, central Laos (Fig. 7). Etymology. The specific epithet ��� lao ��� is derived from both the official name of Laos, the Lao People���s Democratic Republic in which the species was discovered, and the Lao, the main group of people inhabiting Laos., Published as part of Nguyen, Thuong Huyen, Sitthivong, Saly, Ngo, Hanh Thi, Luu, Vinh Quang, Nguyen, Truong Quang, Le, Minh Duc & Ziegler, Thomas, 2020, A new species of Dixonius (Squamata: Gekkonidae) from the karst forest of Khammouane Province, central Laos, pp. 530-542 in Zootaxa 4759 (4) on pages 532-538, DOI: 10.11646/zootaxa.4759.4.4, http://zenodo.org/record/3741047

Ziegler, Thomas, Pham, Cuong The, Nguyen, Tan Van, Nguyen, Truong Quang, Wang, Jian, Wang, Ying-Yong, Stuart, Bryan L., Le, Minh Duc (2019): A new species of Opisthotropis from northern Vietnam previously misidentified as the Yellow-spotted Mountain Stream Keelback O. maculosa Stuart & Chuaynkern, 2007 (Squamata: Natricidae). Zootaxa 4613 (3): 579-586, DOI: https://doi.org/10.11646/zootaxa.4613.3.9

Achalinus timi sp. nov. (Figs. 6–7) Holotype. IEBR A.2018.10 (Field no. PA.180), adult male, from forest within the Copia Nature Reserve, near Hua Ty Village (21°20.105’N, 103°35.860’E), Co Ma Commune, Thuan Chau District, Son La Province, northern Vietnam, collected by A.V. Pham on 12 May 2014 at an elevation of ca. 1470 m above sea level. Diagnosis. A species of the genus Achalinus, characterized by a combination of the following characters: 1) maxillary teeth 27; 2) suture between the internasals distinctly longer than that between the prefrontals; 3) loreal fused with prefrontal on each side, with the prefrontal extending towards the supralabials; 4) dorsal scales in 25–25–23 rows, keeled; 5) ventrals 170+1; 6) subcaudals 72, unpaired; 7) a total length of at least 177.9 mm in males (with a tail length of 37.9 mm and a tail/total length ratio of 0.21); 8) dorsum in preservative reddish to greyish brown above, with wide portion of the vertebral region being distinctly darker; the lower head sides are somewhat paler; infralabial and chin shields light greyish brown; venter greyish cream, with the underside of the tail being somewhat darker and the chin region somewhat paler. Description of holotype. Total length 177.9 mm (SVL 140 mm, TaL 37.9 mm); tail long, tail/total length ratio 0.21; body slender, cylindrical; head length 7.8 mm (from tip of snout to posterior margin of parietal); head slightly distinct from neck, dorsally covered with large shields; eye small, with vertically subelliptic pupil; left maxilla with at least 27 equally sized and curved teeth. Rostral small, triangular, not visible from above; suture between the internasals (1.7 mm) longer than that between the prefrontals (0.8 mm); nostril in the anterior part of the nasal; frontal pentagonal, slightly broader than long, pointed backwards, much shorter than the parietals; parietals bordered each by an elongated nuchal, nuchals separated from each other behind parietals by two small scales; second pair of nuchals about half the size than first pair; loreal lacking, prefrontals stretch towards the supralabials; one supraocular; two anterior temporals, only the smaller, upper one in contact with eye, and two elongated posterior temporals; supralabials six, the first smallest, fourth and fifth in contact with the eye, sixth longest; one mental, followed by six infralabials; first pair of infralabials in contact with each other; first three infralabials in contact with the first pair of chin shields; posterior pair of chin shields smaller, laterally in contact with fourth and fifth infralabials. Dorsal scales elliptical, keeled from the neck region onwards, in 25 rows at the anterior part of the body (one head length behind head), 25 scale rows at midbody, the outer row enlarged, and 23 rows at posterior body (one head length before vent); ventrals 170 (plus one preventral) distinct, laterally rounded; subcaudals 72, unpaired; cloacal entire. The ethanol-preserved holotype is reddish to greyish brown above, with a wide portion of the vertebral region being distinctly darker; the lower head sides somewhat paler; venter greyish cream, with the underside of the tail being somewhat darker and the chin region somewhat paler; infralabial and chin shields light greyish brown. Comparisons. Achalinus timi sp. nov. lacks a separate loreal scale, which is fused with the prefrontal on each side, with the prefrontal extending towards the supralabials; all remaining Achalinus species known at time to occur in Vietnam (A. ater, A. juliani, A. rufescens, and A. spinalis) have a loreal being separated from the prefrontal by a suture, which is also the case in most other Achalinus species (A. hainanus, A. meiguensis, A. niger, and A. werneri) which thus can be easily distinguished from Achalinus timi sp. nov. Achalinus timi sp. nov. further differs from A. formosanus, A. jinggangensis, A. meiguensis, A. rufescens, and A. spinalis by having more maxillary teeth. For details and further distinguishing characters see Table 5. Besides different maxillary teeth counts, Achalinus timi sp. nov. can be distinguished from the two subspecies of A. formosanus as follows: from A. f. formosanus by having the internasal suture distinctly longer than prefrontal suture (versus internasal suture almost as long as prefrontal suture) and 25–25–23 versus 29–27–25 dorsal scale rows; and from A. f. chigirai by 23 versus 25 dorsal scale rows at posterior body and by having more ventrals and less subcaudals (170 versus 161–167 ventrals; 72 versus 96–97 subcaudals). Furthermore, Achalinus timi sp. nov. can be distinguished from A. jinggangensis by having 25 versus 23 dorsal scale rows at midbody, more ventrals and subcaudals (170 versus 156–164 ventrals; 72 versus 51–64 subcaudals, respectively). Etymology. We name this species after Tim N. Ziegler. As common name we propose Tim’s Burrowing Snake. Distribution. The new species currently is only known from the type locality in Son La Province, northwestern Vietnam (Fig. 12). Natural history. The male holotype of Achalinus timi sp. nov. was discovered at 10: 15 in the morning, as a road kill specimen. The surrounding habitat was secondary evergreen forest. Air temperature at the site was 25–30 o C and relative humidity 70–75%., Published as part of Ziegler, Thomas, Nguyen, Truong Quang, Pham, Cuong The, Nguyen, Tao Thien, Pham, Anh Van, Schingen, Mona Van, Nguyen, Tham Thi & Le, Minh Duc, 2019, Three new species of the snake genus Achalinus from Vietnam (Squamata: Xenodermatidae), pp. 249-269 in Zootaxa 4590 (2) on pages 258-260, DOI: 10.11646/zootaxa.4590.2.3, http://zenodo.org/record/2651878

Achalinus juliani sp. nov. (Figs. 2–5) Holotype. IEBR A.2018.8 (Field no. CB 2014.18), an adult male, from forest of Duc Quang Commune (22°43.084’N, 106°39.653’E, at an elevation of 477 m above sea level), Ha Lang District, Cao Bang Province, northern Vietnam, collected by C.T. Pham on 14 June 2014. Paratypes. IEBR A.2018.9 (Field no. CB 2014.19), a female, from Phia Oac-Phia Den National Park (22°36.147’N, 105°52.726’E, at an elevation of 1461 m above sea level), Nguyen Binh District, Cao Bang Province, northern Vietnam, collected by T.H. Pham on 24 May 2014; VNMN 0 6924 (Field no. TAO 766), an adult male, from forest of Thanh Cong Commune within Phia Oac-Phia Den National Park (22°60.556’N, 105°87.530’E, at an elevation of 1590 m above sea level), Nguyen Binh District, Cao Bang Province, northern Vietnam, collected by T.T. Nguyen on 10 May 2010. Diagnosis. A species of the genus Achalinus, characterized by a combination of the following characters: 1) maxillary teeth 28; 2) suture between internasals distinctly longer than that between the prefrontals; 3) internasal not fused to prefrontal; 4) loreal not fused with prefrontal; 5) infralabials 6; 6) mental separated from anterior chin shields; 7) two elongated anterior temporals, in contact with eye, and two posterior temporals; 8) dorsal scales in 25–23–23 rows, keeled; 9) ventrals 173–179; 10) subcaudals 77–91, unpaired; 11) a total length of at least 413 mm (with a maximum tail length of 109 mm, and a tail/total length ratio of 0.22–0.37); 12) dorsum in preservative reddish to greyish brown above, the lower and posterior head sides paler; venter greyish cream, with the underside of the tail being somewhat darker. Description of holotype. Total length 355 mm (SVL 260 mm and TaL 95 mm); tail long, tail/total length ratio 0.365; body slender, cylindrical; head length 8.9 mm (from tip of snout to posterior margin of parietal); head slightly distinct from neck, dorsally covered with large shields; eye small, with vertically subelliptic pupil; left maxilla with 28 equally sized and curved teeth. Rostral small, triangular, slightly visible from above; suture between the internasals (1.4 mm) longer than that between the prefrontals (0.9 mm); nostril in the anterior part of the nasal; frontal pentagonal, slightly broader than long, pointed backwards, much shorter than the parietals; each parietal bordered by an elongated nuchal; nuchals separated from each other behind parietals by two small scales; second pair of nuchals about half the size than first pair; one loreal, distinctly wider than high, extending from the nasal to the eye; one supraocular; two anterior temporals, elongated, upper one smaller, both in contact with eye, and two elongated posterior temporals; supralabials six, the first smallest, fourth and fifth in contact with the eye, sixth longest; third and fourth in contact with the loreal; one mental, followed by six infralabials; first pair of infralabials in contact with each other; first four infralabials in contact with the first pair of chin shields; posterior pair of chin shields smaller, laterally in contact with fourth and fifth infralabials. Dorsal scales elliptical, keeled from the neck region onwards, 23 scale rows at midbody, those of the outer row enlarged, 25 scales round the anterior part of the body (one head length behind head), and 23 dorsal scale rows at posterior body (one head length before vent); ventrals 169 (plus one preventral), distinctly rounded laterally; subcaudals 91, unpaired, plus tail tip; cloacal entire. The ethanol-preserved holotype greyish brown above, the lower and posterior head sides being paler; venter greyish cream, with the underside of the tail being somewhat darker and the chin region somewhat paler; infralabial and chin shields light greyish brown. Ventrals and subcaudals anteriorly and laterally darker. Variation. For scalation details of the paratypes see Table 3. In addition, the nuchals of the female paratype IEBR A.2018.9 are separated from each other behind the parietals by four small scales and those of the male paratype VNMN 0 6924 by two large, two medium sized and a small scale. Furthermore, in the paratype VNMN 0 6924 the posterior pair of chin shields on the left side is in contact with third and fourth infralabials laterally. The hemipenes are basally everted in the specimen VNMN 0 6924. Comparisons. Achalinus juliani sp. nov. can be differentiated from the remaining Achalinus representatives from Vietnam by having more subcaudals, more dorsal scale rows on anterior part of the body, more ventrals, more infralabials, and a longer internasal suture (see Tables 3, 5). Achalinus juliani sp. nov. differs from A. ater in having more dorsal scale rows in the anterior part of the body (25 versus 21–23) and more subcaudals (77–91 versus 47–70). Achalinus juliani sp. nov. differs from A. rufescens by having more infralabials (6 versus 5), more ventrals (163–179 versus 131–158), more subcaudals (77–91 versus 54–82), and more maxillary teeth. Achalinus juliani sp. nov. differs from A. spinalis by the internasal suture being distinctly longer than that between the prefrontals (versus as long as or shorter), more dorsal scale rows in the anterior part of the body (25 versus 21–24), more subcaudals (77–91 versus 39–67), by lacking a black mid-dorsal line, and by having more maxillary teeth. Achalinus juliani sp. nov. differs from A. formosanus by having the internasal suture distinctly longer than prefrontal suture (versus internasal suture almost as long as prefrontal suture), by the loreal not being fused with prefrontal, fewer dorsal scale rows (25–23–23 versus 25–29 – 25–27–25), more maxillary teeth (28 versus 14–17), and by lacking a black mid-dorsal line; in addition, Achalinus juliani sp. nov. can be distinguished from A. f. formosanus by having more subcaudals (77–91 versus 61–83), and from A. f. chigirai by having fewer subcaudals (77–91 versus 96–97). Achalinus juliani sp. nov. differs from A. hainanus by having 6 versus 5 infralabials, 2 versus 1 anterior temporal, more dorsal scale rows in the anterior part of the body (25 versus 23), and more subcaudals (77–91 versus 67–69). Achalinus juliani sp. nov. differs from A. jinggangensis by loreal not being fused with prefrontal, having more ventrals (163–179 versus 156–164), more subcaudals (77–91 versus 51–64), and more maxillary teeth. Achalinus juliani sp. nov. differs from A. meiguensis by mental being separated from anterior chin shields, internasal not fused to prefrontal, by different dorsal scale row counts (25–23–23 versus 21–23 – 19–23 – 19), by having more subcaudals (77–91 versus 39–62), and by having more maxillary teeth. Achalinus juliani sp. nov. differs from A. niger by the internasal suture being distinctly longer than that between the prefrontals (versus almost as long as or shorter), fewer midbody dorsal scale rows (23 versus 25), more subcaudals (77–91 versus 52–72), and keeled body scales (versus smooth on anterior part of body). Achalinus juliani sp. nov. differs from A. werneri by the internasal suture being distinctly longer than that between the prefrontals (versus almost as long as), and by lacking a black mid-dorsal line and a dark subcaudal streak. Etymology. We name this species after Julian L. Ziegler. As common name we propose Julian’s Burrowing Snake. Distribution. The new species currently is only known from two localities within Cao Bang Province, northeastern Vietnam (Fig. 12). Natural history. Specimens were found between 20:00 and 22:00 h on forest paths. The surrounding habitat was secondary forest of large, medium and small hardwoods mixed with shrubs and vines. Air temperature at the sites were 25.4–32.6 o C and relative humidity 65–80%. The new species was found in different habitat types: lowland forest in Phia Oac-Phia Den National Park and limestone karst forest in Ha Lang District, at elevations from 470 to 1460 m above sea level., Published as part of Ziegler, Thomas, Nguyen, Truong Quang, Pham, Cuong The, Nguyen, Tao Thien, Pham, Anh Van, Schingen, Mona Van, Nguyen, Tham Thi & Le, Minh Duc, 2019, Three new species of the snake genus Achalinus from Vietnam (Squamata: Xenodermatidae), pp. 249-269 in Zootaxa 4590 (2) on pages 253-255, DOI: 10.11646/zootaxa.4590.2.3, http://zenodo.org/record/2651878

Lycodon namdongensis sp. nov. Figs. 2 ¯4 Holotype: VNUF R.2017.23 (field number: TT 17.23), adult male, collected on 13 June 2017 by Vinh Quang Luu, Nghia Van Ha, Oanh Van Lo, and Ngoan Van Ha in the karst forest, Nam Dong Nature Reserve, Quan Son District, Thanh Hoa Province, Vietnam (20°18.298’N; 104°54.776’E, at an elevation of 616 m a.s.l.) Diagnosis: Lycodon namdongensis sp. nov is characterized by the following morphological characters: dorsal scales in 17¯17¯15 rows, all smooth; supralabials 8; infralabials 10; one large loreal on each side, separated from the eye; cloacal plate single; ventral scales 218 (plus two preventral scales); subcaudals 85; a total length of 723 mm in the single male; tail / total length ratio 0.205; dorsal head pattern whitish grey; dorsal surface of body with 23 narrowed greyish cream bands; dorsal surface of tail with 14 cream bands; ventral surface of body mostly cream with a few dark spots posteriorly; ventral surface of tail dark grey. Description of the holotype: Head elongate (HL 20.4 mm), moderately distinct from the neck, longer than wide (HW/HL ratio 0.54), depressed (HH/HL ratio 0.40), narrow anteriorly (IN/IO ratio 0.68); snout elongate (SnL/HL ratio 0.33); nostril lateral, oval shaped, located in the middle of the nasal; eye large (ED/HL ratio 0.13), pupils vertically elliptic; rostral triangular, much broader than high, hardly visible from above; nasal divided into two scales by a vertical ridge along posterior edge of nostril; two square internasals, as wide as long, bordered by two large, subpentagonal prefrontals posteriorly; frontal single, enlarged, pentagonal, narrow posteriorly; parietals longer than wide, in contact with each other medially, with upper anterior and posterior temporals, paraparietal laterally and three nuchal scales posteriorly; loreal 1/1 (on the left side the normal sized loreal posteriorly is followed by an additional, very tiny one), elongate, not entering orbit; supralabials 8/8, first and second in contact with nasal, only fifth entering orbit on left side, and third to fifth entering orbit on right side, sixth largest; infralabials 10/10, first pair in broad contact with each other, first to fifth in contact with anterior chin shields; anterior and posterior pairs of chin shields elongate, of about the same size and shape, separated by a medial groove, anterior pair slightly larger than the posterior; preoculars 3/1 (the lower two, tiny preoculars on the left side represent separated upper parts of supralabials 3¯4); postoculars 2/2; anterior temporals 2/2, posterior temporals 2/ 2, upper anterior ones distinctly thinner than lower ones. Left maxilla arched, with an angulous apex, distinctly bent inwards anteriorly. A total of 12 maxillary teeth or teeth alveola, with the following formula: four small anterior teeth, with the last two ones being somewhat enlarged + two strongly enlarged teeth, thick, and not much curved + a wide gap, of about the length of the largest teeth before + three small teeth + a small gap + three posterior teeth, with the first two of them being enlarged. Body elongate, SVL 575 mm; TaL 148 mm; TL 723 mm; ratio of tail length/total length 0.205; ventrals 218 (plus two preventrals); subcaudals 85, divided, weakly notched laterally; cloacal single; DSR 17¯17¯15; all dorsal scale rows smooth; the vertebral scales not enlarged (Table 1). Colouration in life: Head pattern whitish grey; dorsal surface of the head dark in the anterior half and slightly cream posteriorly; ventral side of the head cream, dark-grey anteriorly; dorsal surface of body dark grey with 23 narrowed greyish cream dorsal bands, each band with dark flank spots; first body band starting at ventral scale 25, one vertebral scale covered by this band, and slightly extended laterally to a width of 2–4 (mostly 2) scales; ventral surface mostly cream with a few small dark spots posteriorly; dorsal surface of tail with 14 cream bands, ventral surface of tail dark grey with irregular blotches corresponding to light crossbands, however, these bands do not form complete rings. Hemipenis: The left hemipenis is only in part everted, hemipenis cylindrical at base, lower part smooth, upper part densely covered with spines, slightly widened. Comparisons: Lycodon namdongensis sp. nov. can be distinguished from its closest relative L. futsingensis by having more ventral scales (218 versus 193–208), ventral pattern mostly cream with a few small dark spots posteriorly (versus speckled posteriorly), first body band starting at ventral scale 25 (versus 13–21), and first body band covered by one vertebral scale (versus 5–10) (Vogel et al. 2009; Luu et al. 2013a; Neang et al. 2014), in concert with an uncorrected p -distance of 6.5%; from L. flavozonatus by having cloacal plate single (versus divided), dorsal scale rows being completely smooth (versus 10–12 keeled dorsal scale rows at midbody), a distinctly lower body band count (23 versus 51–78), belly pattern mostly cream with a few small dark spots (versus yellow with large black spots) (Orlov & Ryabov 2004); from L. meridionalis by having whitish grey head pattern (versus with yellow-black marble markings), fewer ventral scales (218 versus 234–246), dorsal scale rows being completely smooth (versus distinctly keeled), dorsal pattern banded (versus blotched), fewer subcaudal scales (86 versus 98–106), ventral surface mostly cream with a few dark spots posteriorly (versus yellow with dark spots posteriorly), and fewer tail bands (14 versus 25–35) (Bourret 1935; Orlov & Ryabov 2004; Gawor et al. 2016); from L. banksi by having a loreal not entering the orbit (versus entering the orbit), fewer ventral scales (218 versus 241), dorsal scale rows being completely smooth (versus feebly keeled), dorsal pattern banded (versus blotched) (Luu et al. 2018); from L. rufozonatus by having a distinctly lower body band count (23 versus 44–52), having fewer dorsal scale rows at neck (17 versus 19), whitish grey dorsal head pattern (versus dark brown with yellow borders), and body bands cream with a few small dark spots laterally (versus yellow or cream, speckled posteriorly) (Boulenger 1893; Deuve 1970); from L. semicarinatus by dorsal scale rows being completely smooth (versus keeled along anterior half), whitish grey dorsal head pattern (versus black), first and second labials dark grey (versus yellow with black sutures), and belly pattern mostly cream with a few small dark spots posteriorly (versus yellow) (Boulenger 1893); from L. capucinus by having more ventrals (218 versus 182–211), fewer supralabials (8 versus 9–10), banded dorsal pattern (versus reticulated), cloacal single (versus divided), and more subcaudal scales (86 versus 59–74) (Neang et al. 2014); from L. cardamomensis by having more light bands on dorsum (23 versus 12–14), fewer dorsal scale rows at neck (17 versus 19), dorsal scale rows being completely smooth (versus keeled on midbody), and more tail bands (14 versus 6–7) (Neang et al. 2014; Do et al. 2017); from L. davidi by having dorsal scale rows being completely smooth (versus middorsal scale rows weakly keeled), slightly fewer ventrals (218 versus 224), body cross-bands greyish cream with dark spots laterally (versus pale tan brown) (Vogel et al. 2012); from L. laoensis by having more ventrals (218 versus 169–192), more subcaudals (85 versus 60–76), fewer supralabials (8 versus 9–10), anterior chinshields slightly longer than posterior ones (versus anterior chinshields 2 to 3 times longer than posterior ones), cloacal single (versus divided) (Vogel & David 2010; Neang et al. 2014); from L. rosozonatum by having fewer dorsal scale rows at neck and at midbody (17–17 versus 19–19, respectively), the vertebral scale row not enlarged (versus distinctly enlarged), dorsal scale rows being completely smooth (versus three to nine upper dorsal scale rows plus vertebral row weakly keeled), and fewer dorsal bands (23 versus 28–35) (Hu et al. 1975); from Lycodon ruhstrati abditus by dorsal scale rows being completely smooth (versus three to five dorsal scale rows plus vertebral row distinctly keeled), fewer subcaudals (85 versus 90–103), first body band starting at ventral scale 25 (versus 12–17), and first body band covering one vertebral scale (versus 5–6) (Vogel et al. 2009; Luu et al. 2013a); from L. zoosvictoriae by having more ventral scales (218 versus 213), dorsum banded (versus blotched), dorsal scale rows being completely smooth (versus weakly keeled), dorsal head whitish grey (versus dark brown), dorsal tail banded (versus blotched) (Neang et al. 2014); from L. multifasciatus by having fewer ventral scales (218 versus 229–237), fewer subcaudals (85 versus 106–119 in males), dorsal scale rows being completely smooth (versus upper dorsal and vertebral rows keeled), tail bands (14 versus 25–42), first body band starting at ventral scale 25 (versus 7–11), and first body band covered by one vertebral scale (versus about 6) (Maki 1931; Vogel et al. 2012); from L. ophiophagus by having more ventral scales (218 versus 211–212), fewer subcaudals (85 versus 87–90), and first body band covered by one vertebral scale (versus about 8) (Vogel et al. 2009; Vogel et al. 2012); from L. paucifasciatus by having fewer dorsal scale rows at neck (17 versus 19), dorsal scale rows being completely smooth (versus upper rows plus vertebral row keeled), fewer subcaudals (85 versus 92 in males), first body band starting at ventral scale 25 (versus 10–15), first body band covered by one vertebral scale (versus 9.5–11), and more light tail bands (14 versus 8–11) (Smith 1943; Vogel et al. 2009; Vogel et al. 2012; Neang et al. 2014); from L. synaptor by having more ventral scale rows (218 versus 201–203), fewer light bands on dorsum (23 versus 30–31), more subcaudals (85 versus 68–69), and dorsal scale rows being completely smooth (versus 6–7 dorsal scale rows keeled) (Vogel & David 2010) (for more details see Table 1). The new species has a loreal which is not in contact with the orbit and thus differs from the following species of the Lycodon fasciatus group which have the loreal entering the orbit: L. butleri Boulenger; L. fasciatus; L. subcinctus. In addition, the new species differs from L. butleri by having fewer dorsal bands (23 versus 28–37), fewer tail bands (14 versus 17–23), belly pattern mostly cream (versus banded & spotted) (Grismer et al. 2014); from L. fasciatus by having dorsal scale rows being completely smooth (versus weakly keeled), dorsal head pattern whitish grey (versus dark), first body band starting at ventral scale 25 (versus 5–18), first body band covered by one vertebral scale (versus 3–12), ventral pattern mostly cream (versus banded) (Vogel & David 2010; Neang et al. 2014); from L. subcinctus by the presence of a preocular scale (versus absent), having dorsal scale rows being completely smooth (versus weakly keeled), single cloacal (versus divided), and more body bands (23 versus 9–15) (Neang et al. 2014) (for more details see Table 1). Distribution: Lycodon namdongensis sp. nov. is currently known only from the type locality in the Nam Dong NR, Thanh Hoa Province, Vietnam. Distribution of the new species and other Lycodon species from Vietnam was shown in Figs. 5 and 6. Etymology: We name this species after its type locality, Nam Dong NR, where the new Lycodon was discovered and propose the following common names: Namdong Wolf Snake (English), Namdong Wolfszahnnatter (German), Rǻn khuyḗt nam động (Vietnamese). Natural history: The holotype was collected at 7:54 PM, while crawling on the forest path, at an elevation of 616 m a.s.l. The surrounding habitat was shrubs in secondary karst forest (Fig. 7). The humidity at the time of collection was approximately 66% and the air temperature was 28.4 o C.

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

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

Cyrtodactylus taybacensis sp. nov. (Figs. 3���5) Holotype. IEBR 4379 (Field No. SL 2016.400), adult male, collected by A.V. Pham and D.A. Giang on 14 October 2016 in the karst forest near Ca Nang Village (21˚54.440���N, 103˚31.390���E, elevation: 650 m asl.), Ca Nang Commune, Quynh Nhai District, Son La Province, northwestern Vietnam. Paratypes. TBU 0 7, 0 8 (Field Nos. SL 2016.271, 272), adult males collected by A.V. Pham, T. Q.L. Hoang, L. M. Ha, N. B. Song, D. K.K.S. Vanh, and C. A. Lau on 25 September 2016 and IEBR 4380 (Field No. SL 2017.26), adult female, collected by N.B. Song and D. K.K.S. Vanh on 22 June 2017 in the karst forest near Pha Luong Village (21˚36.324���N, 103˚34.540���E, elevation: 720 m asl.), Phong Lai Commune, Thuan Chau District, Son La Province; IEBR 4381 & 4382 (Field Nos. SL2016.403, 405), adult males and TBU 0 9 (Field No. SL 2016.404), adult female collected by A.V. Pham, L. M. Ha, O. V. Dieu and D. A. Giang on 15 October 2016 near Tua Thang Commune, Tua Chua District, Dien Bien Province (21˚56.933���N, 103˚27.803���E, elevation: 610 m asl.). Diagnosis. The new species can be distinguished from other members of the genus Cyrtodactylus by a combination of the following characters: medium size (SVL up to 97.5 mm); dorsal tubercles in 13���16 irregular rows; ventral scale rows 30���38; ventrolateral folds present without interspersed tubercles; each thigh with 11���13 enlarged femoral scales; femoral pores absent in males and females; precloacal pores 11���13 in males, five or 15 pitted scales in females, in a continuous row; postcloacal tubercles two or three; lamellae under toe IV 16���20; subcaudal scales transversely enlarged; dorsal head with dark brown marking, oval, triangle and arched shape; five brown dorsal bands between limb insertions. Description of holotype. Adult male, snout-vent length (SVL) 85.7 mm; body elongate (AG/SVL 0.46); head distinct from neck, elongate, depressed (HL/SVL 0.28, HW/HL 0.67, HH/HL 0.41); supranasals in contact with each other anteriorly, separated from each other by a small scale posteriorly; nares oval, surrounded by supranasal, rostral, first supralabial, and three postnasals; loreal region concave; snout long (SE/HL 0.38), round anteriorly, longer than diameter of orbit (OD/SE 0.68); snout scales small, round, granular, larger than those in frontal and parietal regions; eye large (OD/HL 0.26), pupils vertical; upper eyelid fringe with spinous scales; ear opening oval, obliquely directed, small in size (ED/HL 0.07); rostral wider than high (RH/RW 0.74), bordered by first supralabial, nostril and supranasal on each side; mental triangular, as wide as rostral (RW 3.6 mm, MW 3.5 mm), wider than high (ML/MW 0.74); postmentals two, enlarged, in contact with mental anteriorly, first infralabial laterally, and eight small scales posteriorly; supralabials 10/11; infralabials 9/9. Dorsal scales granular; dorsal tubercles round, three or four times larger than the size of adjoining scales, conical, present on occiput, back and tail base, each surrounded by 9 or 10 granular scales, in 15 or 16 irregular longitudinal rows at midbody; ventral scales smooth, medial scales two times larger than dorsal scales, round, subimbricate, in 34���36 longitudinal rows at midbody; lateral skin folds distinct, without tubercles; gular region with homogeneous smooth scales; ventral scales between mental and cloacal slit 177; precloacal groove absent; three rows of enlarged scales present in posterior region of pore-bearing scales; enlarged femoral scales present; femoral pores absent; precloacal pores 13, in a continuous row. Fore and hind limbs moderately slender (ForeaL/SVL 0.18, CrusL/SVL 0.23); dorsal surface of forelimbs covered by small scales, slightly larger than underside scales, without tubercles; dorsal surface of hind limbs covered by distinctly developed tubercles; interdigital webbing weakly developed; subdigital lamellae: finger I 14/ 14 (with 6/6 basally broadened lamellae), finger II 16/15 (7/6), finger III 17/18 (6/6), finger IV 18/18 (7/7), finger V 16/17 (6/5), toe I 15/15 (6/6), toe II 15/16 (6/6), toe III 17/17 (6/6), toe IV 20/19 (9/8), toe V 19/20 (7/7). Tail regenerated; postcloacal tubercles 2/2; dorsal tail base with distinct tubercles; subcaudals distinctly enlarged in the original part, smooth. ������continued on the next page Coloration in life. Ground color of dorsal head and back yellowish brown; snout region yellow with two dark spots; dorsal head with dark marking, oval, triangle and arched shape; a dark stripe extending from posterior corner of eye to above tympanum; labials yellow; neck with some large dark blotches, forming a discontinuous band anteriorly and a continuous band posteriorly; dorsum with five transverse dark brown bands between fore and hind limb insertions, edged in yellow anteriorly and posteriorly; dorsal surface of fore and hind limbs with dark brown blotches; dorsal surface of tail yelowish brown with two dark bands at base, regenerated part greyish cream; chin, throat, chest, belly and lower limbs pinkish white; ventral surface of tail greyish (Fig. 3). Sexual dimorphism and variation. The females differ from male specimens in the absence of hemipenial swellings at the tail base and having a larger size. The males have 11���13 precloacal pores (versus 5 or 15 pitted scales in females). The tails of paratypes (TBU 0 7 and IEBR 4380) are complete, greyish cream with nine dark brown bands, subcaudals distinctly enlarged approximately three fourths of tail length. For other morphological characters see Table 1. Distribution. Cyrtodactylus taybacensis sp. nov. is currently known from Tua Chua District in Dien Bien Province and Thuan Chau and Quynh Nhai districts in Son La Province, Vietnam (Fig. 1). Etymology. Specific epithet ��� taybacensis ��� is a toponym in reference to the type locality of the species, meaning ���from northwestern region���. For the common names we suggest Taybac Bent-toed Gecko (English) and Th���ch s��ng ng��n t��y b��c (Vietnamese). Natural history. Specimens were found at night between 19:00 and 21:00, on trees near limestone cliffs and in rock crevices, approximately 0.5���1.5 m above the ground, at elevations between 600 and 720 m asl. The surrounding habitat was disturbed evergreen karst forest of medium hardwood and shrub. The relative humidity was approximately 75���85% and the air temperature ranged from 25 to 34 o C. Comparisons. We compared the new species with its congeners from Vietnam and neighboring countries in mainland Indochina, including Laos, Cambodia, Myanmar, China (Yunnan), Thailand, and peninsular Malaysia based on examination of specimens (see Appendix) and data obtained from the literature (Smith 1917, 1921a, b, 1935; Taylor 1963; Ulber & Grossmann 1991; Ulber 1993; Bauer 2002, 2003; Bauer et al. 2002, 2003, 2009, 2010; Ziegler et al. 2002, 2010, 2013; Pauwels & Sumontha 2014; Pauwels et al. 2004, 2013, 2014a, b, 2016; Nguyen et al. 2006, 2014; Hoang et al. 2007; Orlov et al. 2007; Grismer & Ahmad 2008; Ngo 2008, 2011; Ngo & Bauer 2008; Ngo & Chan 2010, 2011; Ngo & Grismer 2010, 2012; Ngo & Pauwels 2010; Ngo et al. 2008, 2010; Sumontha et al. 2010, 2012, 2014; Chan-ard & Makchai 2011; David et al. 2011; Schneider et al. 2011; Luu et al. 2011, 2014, 2015, 2016a, b, c; Grismer et al. 2012, 2016, 2017; Kunya et al. 2014, 2015; Nazarov et al. 2014; Panitvong et al. 2014; Le et al. 2016; Connette et al. 2017; Pham et al. 2017; and Nguyen et al. 2015a, 2017). The new species can be distinguished from other known species of Cyrtodactylus by morphological characters (see Table 2). Below we compared the new species from the closely related species from northern Vietnam and northern Laos. In general appearance, Cyrtodactylus taybacensis sp. nov. is similar to C. bichnganae Ngo & Grismer, C. cf. bichnganae, C. bobrovi Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, C. cucphuongensis Ngo & Chan, C. huongsonensis Luu, Nguyen, Do & Ziegler, C. martini Ngo, C. puhuensis Nguyen, Yang, Le, Nguyen, Orlov, Hoang, Nguyen, Jin, Rao, Hoang, Che, Murphy & Zhang, C. otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, C. spelaeus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov, C. soni Le, Nguyen, Le & Ziegler, C. sonlaensis Nguyen, Pham, Ziegler, Ngo & Le, C. vilaphongi Schneider, Nguyen, Duc Le, Nophaseud, Bonkowski & Ziegler, C. wayakonei Nguyen, Kingsada, R��sler, Auer & Ziegler. Cyrtodactylus taybacensis sp. nov. differs from C. bichnganae by having more precloacal pores in males (11���13 vs. 10) and the absence of femoral pores on each thigh in males (vs. 9); from C. bobrovi by having fewer ventral scale rows (30���38 vs. 40���45), the presence of enlarged femoral scales (vs. absent), the presence of enlarged subcaudals (vs. absent), more preloacal pores in males (11���13 vs. 5), and fewer lamellae under toe IV (16���20 vs. 21���22); from C. chauquangensis by the presence of enlarged femoral scales (vs. absent) and having more precloacal pores in males (11���13 vs. 6); from C. cucphuongensis by the presence of precloacal pores in males (vs. absent), having fewer ventral scale rows (30���38 vs. 42), fewer lamellae under finger IV (17���19 vs. 21) and under toe IV (16���20 vs. 24); from C. huongsonensis by having more enlarged femoral scales on each thigh (11���14 vs. 7���9), more precloacal pores in males (11���13 vs. 6), and fewer ventral scale rows (30���38 vs. 41���48); from C. martini by having more precloacal pores in males (11���13 vs. 4), fewer ventral scale rows (30���38 vs. 39���43), and fewer lamellae under toe IV (16���20 vs. 22���24); from C. puhuensis by having more precloacal pores in males (11���13 vs. 5) and fewer lamellae under toe IV (16���20 vs. 23); from C. otai by having more preloacal pores in males (11���13 vs. 7���8), the presence of enlarged femoral scales (vs. absent), and the presence of enlarged subcaudals (vs. absent); from C. soni by having more enlarged femoral scales on each thigh (11���14 vs. 8���9), more precloacal pores in males (11���13 vs. 6���8), the absence of femoral pores in males (vs. 6���8 on each thigh), and fewer ventral scale rows (30���38 vs. 41���45); from C. sonlaensis by its larger size (SVL reaching 97.5 mm vs. 83.2 mm), the absence of femoral pores in males (vs. 13���15 on each thigh), more precloacal pores in males (11���13 vs. 8), and fewer enlarged femoral scales on each thigh (11���14 vs. 15���17); from C. spelaeus by having more precloacal pores in males (11���13 vs. 8���9) and fewer lamellae under toe IV (16���20 vs. 22���24); from C. vilaphongi by the presence of enlarged subcaudals (vs. absent), the presence of of enlarged femoral scales (vs. absent), and the presence of pitted scales in females (vs. absent); and from C. wayakonei by having fewer dorsal tubercle rows (13���16 vs. 16���19), the presence of enlarged femoral scales (vs. absent), and more precloacal pores in males (11���13 vs. 6���8). Morphologically, Cyrtodactylus taybacensis sp. nov. resembles C. cf. bichnganae. However, the new species can be distinguished from the latter by having more precloacal pores in males (11���13 vs. 7���9 in C. cf. bichnganae) and the absence of femoral pores on each thigh in males (vs. 3���10 in C. cf. bichnganae)., Published as part of Pham, Anh Van, Le, Minh Duc, Ngo, Hanh Thi, Ziegler, Thomas & Nguyen, Truong Quang, 2019, A new species of Cyrtodactylus (Squamata: Gekkonidae) from northwestern Vietnam, pp. 360-380 in Zootaxa 4544 (3) on pages 364-369, DOI: 10.11646/zootaxa.4544.3.3, http://zenodo.org/record/2618393, {"references":["Smith, M. A. (1917) Descriptions of new reptiles and a new batrachian from Siam. Journal of Natural History Society of Siam, 2, 221 - 225.","Smith, M. A. (1921 a) New or little-known reptiles and batrachians from southern Annam (Indo-China). Proceedings of the Zoological Society of London, 1921, 423 - 440.","Smith, M. A. (1921 b) Reptiles and Batrachians collected on Pulo Condore. Journal Natural History Society, 4 (1920), 93 ‾ 97.","Smith, M. A. (1935) The fauna of British India, including Ceylon and Burma. Fol. II. Sauria. Taylor and Francis, London, 8 + 440 pp.","Taylor, E. H. (1963) Lizards of Thailand. The University of Kansas Science Bulletin, 55, 687 - 1077.","Ulber, T. & Grossmann, W. (1991) Ein weiterer neuer Gecko aus Zentral-Thailand: Cyrtodactylus papilionoides sp. nov. (Reptilia: Sauria: Gekkonidae). Sauria, 13, 13 - 22.","Ulber, T. (1993) Bemerkungen uber cyrtodactyline Geckos aus Thailand nebst Beschreibungen von zwei neuen Arten (Reptilia: Gekkonidae). Mitteilungen aus dem Zoologischen Museum in Berlin, 69, 187 - 200. https: // doi. org / 10.1002 / mmnz. 19930690202","Ziegler, T., Rosler, H., Herrmann, H. - W. & Vu, T. N. (2002) Cyrtodactylus phongnhakebangensis sp. n., ein neuer Bogenfingergecko aus dem annamitischen Karstwaldmassiv, Vietnam. Herpetofauna, 24, 11 - 25.","Ziegler, T., Nazarov, R., Orlov, N., Nguyen, T. Q., Vu, T. N., Dang, K. N., Dinh, T. H. & Schmitz, A. (2010) A third new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha-Ke Bang National Park, Vietnam. Zootaxa, 2413 (1), 20 - 36. https: // doi. org / 10.11646 / zootaxa. 2413.1.2","Ziegler, T., Phung, T. M., Le, M. D. & Nguyen, T. Q. (2013) A new Cyrtodactylus (Squamata: Gekkonidae) from Phu Yen Province, southern Vietnam. Zootaxa, 3686 (4), 432 - 446. https: // doi. org / 10.11646 / zootaxa. 3686.4.2","Sumontha, M., Pauwels, O. S. G., Suwannakarn, N., Nutatheera, N. & Sodob, W. (2014) Cyrtodactylus wangkulangkulae (Squamata: Gekkonidae), a new Bent-toed Gecko from Satun Province, southern Thailand. Zootaxa, 3821 (1), 116 - 124. https: // doi. org / 10.11646 / zootaxa. 3821.1.8","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.","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. https: // doi. org / 10.11646 / zootaxa. 3635.3.7","Pauwels, O. S. G., Sumontha, M., Keeratikiat, K. & Phanamphon, E. (2014 a) Cyrtodactylus kunyai (Squamata: Gekkonidae), a new cave-dwelling Bent-toed Gecko from Loei Province, northeastern Thailand. Zootaxa, 3821 (2), 253 - 264. https: // doi. org / 10.11646 / zootaxa. 3821.2.5","Pauwels, O. S. G., Sumontha, M., Panitvong, N. & Varaguttanonda, V. (2014 b) Cyrtodactylus khelangensis, a new cave-dwelling Bent-toed Gecko (Squamata: Gekkonidae) from Lampang Province, northern Thailand. Zootaxa, 3755 (6), 584 - 594. https: // doi. org / 10.11646 / zootaxa. 3755.6.5","Pauwels, O. S. G., Sumontha, M. & Bauer, A. M. (2016) A new Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Phetchaburi Province, Thailand. Zootaxa, 4088 (3), 409 - 419. https: // doi. org / 10.11646 / zootaxa. 4088.3.6","Nguyen, S. N., Orlov, N. L. & Darevsky, S. I. (2006) Descriptions of two new species of the genus Cyrtodactylus Gray, 1827 (Squamata: Sauria: Gekkonidae) from southern Vietnam. Russian Journal of Herpetology, 13, 215 - 226.","Hoang, Q. X., Orlov, N. L., Ananjeva, N. B., Johns, A. G., Hoang, T. N. & Dau, V. Q. (2007) Description of a new species of the genus Cyrtodactylus Gray, 1827 (Squamata: Sauria: Gekkonidae) from the karst of North Central Vietnam. Russian Journal of Herpetology, 14, 98 - 106.","Orlov, N. L., Nguyen, Nazarov, R. A., Anajeva, N. B. & Nguyen, S. N. (2007) A new species of the genus Cyrtodactylus Gray, 1827 and redescription of Cyrtodactylus paradoxus (Darevsky et Szczerbak, 1997) [Squamata: Sauria: Gekkonidae] from south Vietnam. Russian Journal of Herpetology, 14, 145 - 152.","Grismer, L. L. & Ahmad, N. (2008) A new insular species of Cyrtodactylus (Squamata: Gekkonidae) from the Langkawi Archipelago, Kedah, Peninsular Malaysia. Zootaxa, 1924, 53 - 68.","Ngo, T. V. & Bauer, A. M. (2008) Descriptions of two new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) endemic to southern Vietnam. Zootaxa, 715, 27 - 42.","Ngo, T. V. & Chan, K. O. (2010) A new species of Cyrtodactylus Gray, 1826 (Squamata: Gekkonidae) from Khanh Hoa province, southern Vietnam. Zootaxa, 2504, 47 - 60.","Ngo, T. V. & Chan, K. O. (2011) A new karstic cave-dwelling Cyrtodactylus Gray (Squamata: Gekkonidae) from northern Vietnam. Zootaxa, 3125, 51 - 63.","Ngo, T. V., Grismer, L. L. & Grismer, J. L. (2010) A new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) in Phu Quoc National Park, Kien Giang Biosphere Reserve, southwestern Vietnam. Zootaxa, 2604, 37 - 51.","Ngo, T. V. & Pauwels, O. S. G. (2010) A new cave-dwelling species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Khammouane Province, southern Laos. Zootaxa, 2730, 44 - 56. https: // doi. org / 10.11646 / zootaxa. 2730.1.3","Sumontha, M., Panitvong, N. & Deein, G. (2010) Cyrtodactylus auribalteatus (Squamata: Gekkonidae), a new cave-dwelling gecko from Phitsanulok Province, Thailand. Zootaxa, 2370 (1), 53 - 64. https: // doi. org / 10.11646 / zootaxa. 2370.1.3","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 (1), 7 - 15.","David, P., Nguyen, T. Q., Schneider, N. & Ziegler, T. (2011) A new species of the genus Cyrtodactylus Gray, 1827 from central Laos (Squamata: Gekkonidae). Zootaxa, 2833, 29 - 40.","Schneider, N., Nguyen, T. Q., Schmitz, A., Kingsada, P., Auer, M. & Ziegler, T. (2011) A new species of karst dwelling Cyrtodactylus (Squamata: Gekkonidae) from northwestern Laos. Zootaxa, 2930, 1 - 21.","Luu, V. Q., Nguyen, T. Q., Do, H. Q. & Ziegler, T. (2011) A new Cyrtodactylus (Squamata: Gekkonidae) from Huong Son limestone forest, Hanoi, northern Vietnam. Zootaxa, 3129, 39 - 50.","Luu, V. Q., Calame, T., Bonkowski, M., Nguyen, T. Q. & Ziegler, T. (2014) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Khammouane Province, Laos. Zootaxa, 3760 (1), 54 ‾ 66. https: // doi. org / 10.11646 / zootaxa. 3760.1.3","Luu, V. Q., Calame, T., Nguyen, T. Q., Bonkowski, M. & Ziegler, T. (2015) A new species of Cyrtodactylus (Squamata: Gekkonidae) from the limestone forest of Khammouane Province, central Laos. Zootaxa, 4058 (3), 388 ‾ 402. https: // doi. org / 10.11646 / zootaxa. 4058.3.6","Luu, V. Q., Bonkowski, M., Nguyen, T. Q., Le, M. D., Schneider, N., Ngo, H. T. & Ziegler, T. (2016 a) Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos. Zootaxa, 4107 (2), 101 ‾ 140. https: // doi. org / 10.11646 / zootaxa. 4107.2.1","Luu, V. Q., Calame, T., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016 b) Cyrtodactylus rufford, a new cavedwelling Bent-toed Gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4067 (2), 185 ‾ 199. https: // doi. org / 10.11646 / zootaxa. 4067.2.4","Luu, V. Q., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016 c) A new species of karst-dwelling Bent-toed Gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4079 (1), 87 ‾ 102. https: // doi. org / 10.11646 / zootaxa. 4079.1.6","Grismer, L. L., Wood, P. L. Jr., Quah, E. S. H., Anuar, S., Muin, 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.","Grismer, L. L., Wood, P. L. Jr., Anuar, S., Davis, H. R., Burch, B. T., Cobos, A. J. & Murdoch, M. L. (2016) A new species of karst forest Bent-toed Gecko (genus Cyrtodactylus Gray) not yet threatened by foreign cement companies and a summary of Peninsular Malaysia's endemic karst forest herpetofauna and the need for its conservation, Zootaxa, 4061 (1), 1 - 17. https: // doi. org / 10.11646 / zootaxa. 4061.1.1","Grismer, L. L, Wood, P. L. Jr., Thura, M. K., Zin, T., Quah, E. S. H., Murdoch, M. L., Grismer, M. S., Lin, A., Kyaw, H. & Lwin, N. (2017) Twelve new species of Cyrtodactylus Gray (Squamata: Gekkonidae) from isolated limestone habitats in eastcentral and southern Myanmar demonstrate high localized diversity and unprecedented microendemism. Zoological Journal of the Linnean Society, 1 - 98. https: // doi. org / 10.1093 / zoolinnean / zlx 057","Kunya, K., Panmongkol, A., Pauwels, O. G. S., Sumontha, M., Meewasana, J., Bunkhwamdi, W. & Dangsri, S. (2014) A new forest-dwelling Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Doi Suthep, Chiang Mai Province, northern Thailand. Zootaxa, 3811 (2), 251 - 261. https: // doi. org / 10.11646 / zootaxa. 3811.2.6","Kunya, K., Sumontha, M., Panitvong, N., Dongkumfu, W., Sirisamphan, T. & Pauwels, O. S. G. (2015) A new forest-dwelling Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Doi Inthanon, Chiang Mai Province, northern Thailand. Zootaxa, 3905 (4), 573 - 584. https: // doi. org / 10.11646 / zootaxa. 3905.4.9","Nazarov, R. A., Poyarkov, N. A., Orlov, N. L., Nguyen, S. N., Milto, K. D., Martynov, A. A., Konstantinov, E. L. & Chulisov, A. S. (2014) A review of genus Cyrtodactylus (Reptilia: Sauria: Gekkonidae) in fauna of Laos with description of four new species. Proceedings of the Zoological Institute RAS, 318, 391 - 423.","Panitvong, N., Sumontha, M., Tunprasert & J. Pauwels, O. S. G. (2014) Cyrtodactylus saiyok sp. nov., a new dry evergreen forest-dwelling Bent-toed Gecko (Squamata: Gekkonidae) from Kanchanaburi Province, western Thailand. Zootaxa, 3869 (1), 64 - 74. https: // doi. org / 10.11646 / zootaxa. 3869.1.6","Le, D. T., Nguyen, T. Q., Le, M. D. & Ziegler, T. (2016) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Ninh Binh Province, Vietnam. Zootaxa, 4162 (2), 268 - 282. https: // doi. org / 10.11646 / zootaxa. 4162.2","Connette, G. M., Oswald, P., Thura, M. K., Connette, K. J. L., Grindley, M. E., Songer, M., Zug, R. G. & Mulcahy, G. D. (2017) Rapid forest clearing in a Myanmar proposed national park threatens two newly discovered species of geckos (Gekkonidae: Cyrtodactylus). PLOS ONE, 12 (4), e 0174432. https: // doi. org / 10.1371 / journal. pone. 0174432","Pham, A. V., Pham, N. V., Nguyen, L. D. T., Le, M. D., Do, Q. H., Pham, N. D. & Nguyen, T. Q. (2017) Additional morphological characters and molecular divergence of Cyrtodactylus bichnganae (Reptilia: Gekkonidae). Proceedings of the Seventh National Scientific Conference on Ecology and Biological Resources, 2017, pp 554 ‾ 561. [in Vietnamese with English abstract]","Nguyen, T. Q., Le, M. D., Pham, A. V., Ngo, H. N., Hoang, C. V., Pham, C. T. & Ziegler, T. (2015 a) Two new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of Hoa Binh Province, Vietnam. Zootaxa, 3985 (3), 375 - 390. https: // doi. org / 10.11646 / zootaxa. 3985.3.3"]}

Parafimbrios vietnamensis sp. nov. Figs. 2–3. Holotype. IEBR A.2018.7 (field number LC2016.8), male, collected by H.V. Tu and H.N. Ngo on 2 nd June 2016 near Hoang Ho Village, Phang So Lin Commune, Sin Ho District, Lai Chau Province (22 0 22.180’N, 103 0 14.485’E), at an elevation of 1,317 m above sea level. Diagnosis. A species of the genus Parafimbrios, characterized by a combination of the following characters: 1) rostral laterally with two raised, curved edges; the upper one, together with a horizontal curved ridge of tissue, separate the rostral from the internasals; 2) nasal in contact zone with rostral with curved raised edge; 3) nasal in contact zone with supralabials each with two small oblique, curved raised edges located above first and second as well as above second and third supralabials; 4) suture between the internasals much longer than that between the prefrontals; 5) supralabials 8, with the first four bearing raised edges; 6) infralabials 7; mental and anterior three to four infralabials with raised edges; 7) temporals 4+4–5; 8) 35–33–29 dorsal scale rows; 9) laterally rounded ventrals 164; 10) unpaired subcaudals 49; 11) total length of at least 266 mm in males (with a tail length of 44 mm, and a TaL/TL ratio of 0.165). Description of holotype. Total length 266 mm (snout-vent length 222 mm, tail length 44 mm); TaL/TL ratio 0.165; body slender, cylindrical; head length 7.2 mm (from tip of snout to hind margin of parietal); head not distinct from neck, dorsally covered with large shields; eye small, with vertically sub-elliptic pupil; right maxilla with ca. 27 small, curved, subequal teeth, progressively slightly enlarged posteriorly. Rostral triangular, not visible from above; the sides of the rostral each with two raised, curved edges; the upper one, together with a horizontal curved ridge of tissue at the outermost front of the internasals separate the rostral from the internasals; suture between the internasals (1.7 mm) much longer than that between the prefrontals (0.7 mm); nostril in the anterior part of a large, concave nasal; nasal in contact zone with rostral with curved raised edge, extending from horizontal, curved ridge of tissue in front of each internasal to first supralabial; nasal in contact zone with supralabials each with two small oblique, curved raised edges located above first and second as well as above second and third supralabials; frontal slightly broader than long, rounded at both sides in front, much shorter than the parietals, from anterior tip subdivided in middle for most part (3/4) of its length; loreal large, heptagonal, extending from the nasal to the eye; one small preocular, at the upper front of eye; one supraocular, wider than high; two postoculars on both sides, the upper one hexagonal, being much larger, the lower one elongated, reaching somewhat below the eye; on the right side a very small third postocular, discernible, reaching somewhat above the eye; one large pentagonal to hexagonal subocular; four anterior and four (left) to five (right) posterior temporals each; eight supralabials, the first five very small, with the first four bearing raised edges, the last one much elongated; sixth and seventh supralabials in contact with the subocular; one mental, followed by seven infralabials; mental and anterior three to four infralabials with their raised edges; first pair of infralabials in contact with each other; first five infralabials in contact with the single pair of enlarged chin shields; chin shields covering nearly the whole of the chin in front, in contact with the first ventral; no posterior chin-shields. Dorsal scales small, cycloid, keeled from region behind the neck onwards; 33 scale rows at midbody, every second of the outermost row distinctly enlarged; ca. 35 scales round the anterior part of the body, and 29 dorsal scale rows at posterior body; correspondence of two dorsal scale rows per ventral plate throughout the body; distinct, laterally rounded ventrals 164; single subcaudals 49, without tail tip; cloacal entire; tail moderate. The left hemipenis basally everted. Coloration of holotype in preservation. The ethanol-preserved holotype brownish-black above, with broad yellow neck band widening towards the venter and stretching to the chin region; dorsal head surface in part reddish-brown, in particular in the middle of parietals and frontal; venter greyish-brown, lighter anteriorly, darker towards tail region; ventrals anteriorly and laterally darker. Etymology. Analogous to the naming of the recent Parafimbrios discovery from Laos (P. lao) we name the new species from Vietnam after the country of origin. Distribution. Parafimbrios vietnamensis sp. nov. currently is only known from the type locality in Lai Chau Province, Vietnam (Fig. 4). Natural history. The male holotype of Parafimbrios vietnamensis sp. nov. was discovered at 21:00 at an elevation of 1,317 m a.s.l. in a forest valley. It was drizzling at the time of capture with the air temperature at the site being 24 ° C and the relative humidity 85%. The surrounding habitat was disturbed limestone karst forest and fragmented with hill agricultural fields.

Lycodon banksi sp. nov. Figs 2-5, Table 3 Holotype: VNUF R.2015.20 (field number: TK 20.15), adult male, collected on 4 April 2015 by Vinh Quang Luu and Thomas Calame in the karst forest, at the mouth of a cave, Phou Hin Poun NPA, Hinboun District, Khammouane Province, central Laos, at an elevation of 167 m a.s.l. Diagnosis: Lycodon banksi sp. nov is characterized by the following morphological characters: (1) dorsal scales in 17-17-15 rows, dorsal scales on the anterior 2/3 of the body length smooth, the six central dorsal scale rows of the posterior 1/3 of the body length feebly keeled; (2) supralabials 8; (3) infralabials 10; (4) loreal entering orbit; (5) cloacal single; (6) ventral scales 241; (7) dorsal surface of body with 87 greyish yellow blotches; (8) ventral surface of body and tail uniformly grey cream. Description of the holotype: Head elongate (HL 15.3 mm), moderately distinct from the neck, longer than wide (HW/HL ratio 0.71), depressed (HH/HL ratio 0.40), narrow anteriorly (IN/IO ratio 0.65); snout elongate (SnL/HL ratio 0.39); nostril lateral, oval shaped, located in the middle of the nasal; eye large (ED/HL ratio 0.17), pupils vertically elliptic; rostral triangular, much broader than high, hardly visible from above; nasal divided into two scales by a vertical ridge along posterior edge of nostril; two square internasals, as wide as long, bordered by two large, subpentagonal prefrontals posteriorly; frontal single, enlarged, pentagonal, narrowed posteriorly; parietals longer than wide, in contact with each other medially, with upper anterior and posterior temporals, paraparietal laterally and four nuchal scales posteriorly; loreal 1/1, elongate, entering orbit; supralabials 8/8, first and second in contact with nasal, third to fifth entering orbit, sixth largest; infralabials 10/10, first pair in broad contact with each other, first to fifth in contact with first pair of chin shields; first and second pairs of chin shields elongate, of the same size and shape, separated by a medial groove, first pair larger than the second; preocular 1/1; postoculars 2/2, of the same size, bordering anterior temporals; anterior temporals 2/2, posterior temporals 3/3, upper ones smaller than lower ones. Body elongate, SVL 415 mm; TaL> 50 mm (tail tip lost); preventrals 2, ventrals 241; subcaudals 26 (tail tip lost), divided, weakly notched laterally; cloacal single; DSR 17-17-15; dorsal scales on the anterior 2/3 of the body length smooth, the six central dorsal scale rows of the posterior 1/3 of the body length feebly keeled; the vertebral scales not enlarged. Colouration in life: Head dark grey, without vertical light nuchal band; dorsal surface of body dark greyyellow with 87 greyish yellow irregular dorsal blotches; first body blotch starting at ventral scale 13, a half vertebral scale covered by this blotch; two yellow stripes on each side, from behind the neck to vent, indistinct posteriorly; ventral scales grey cream; dorsal surface of tail with at least eleven greyish yellow tail blotches, ventral surface of tail grey cream. Hemipenis: The left hemipenis is only in part everted but shows a spinose ornamentation. Additional specimen: One specimen which was not collected but detected and photographed on 22 July 2016 by an arachnology team consisting of Peter Jaeger, Aloke Sahu and Jonas Ewert, in Khammouane Province, in ca. 12.4 km distance from the type locality. The color pattern of this specimen resembles closely that of the holotype. Comparisons: In our phylogenetic analysis, Lycodon banksi sp. nov. is nested in a clade containing L. rufozonatus, L. semicarinatus (Cope), ‘ L. flavozonatus’, L. futsingensis and L. meridionalis. The new species differs from the similar L. meridionalis by having loreal entering the orbit (versus separated from the orbit), dorsal scales on the anterior 2/3 of the body length smooth, the six central dorsal scale rows on the posterior body third feebly keeled (versus distinctly keeled), dorsal head pattern uniform dark grey (versus with yellow-black marbling in L. meridionalis), and ventral surface grey cream (versus yellow with dark spots posteriorly) (see Bourret, 1935; Orlov & Ryabov, 2004); from L. rufozonatus by having loreal entering the orbit (versus usually separated), a distinctly higher ventral scale count (241 versus 185-204), dorsal scales feebly keeled in the posterior body part (versus all smooth), dorsal head pattern uniform dark grey (versus dark brown with yellow borders), and body pattern blotched (versus banded) (Boulenger 1893); from L. semicarinatus by having loreal touching the orbit (versus separated), a higher ventral scale count (241 versus 211-234), dorsal scale rows keeled along posterior 1/3 (versus keeled along anterior half), belly pattern uniform grey cream (versus yellow), and body pattern blotched (versus banded) (Boulenger 1893); from L. flavozonatus by having loreal in contact with the orbit (versus separated), cloacal single (versus divided), six dorsal scale rows on the posterior third of the body feebly keeled (versus 10-12 keeled dorsal scale rows at midbody), dorsal head dark grey (versus black with light markings), and belly pattern uniform grey cream (versus yellow with large black spots); from L. futsingensis by having loreal entering the orbit (versus separated), a higher ventral scale count (241 versus 193-203 in males), dorsal scales feebly keeled in the posterior body part (versus all smooth), and body pattern blotched (versus banded) (Vogel et al., 2012; Neang et al., 2014) (Table 4). The new species has a loreal entering the orbit and thus differs from the following species and subspecies of the Lycodon ruhstrati group which have the loreal separated from the orbit: L. cardamomensis Daltry & Wüster, 2002, L. davidi, L. multifasciatus (Maki, 1931), L. ophiophagus Vogel, David, Pauwels, Sumontha, Norval, Hendrix, Vu & Ziegler, 2009, L. paucifasciatus Rendahl in Smith, 1943, L. ruhstrati ruhstrati (Fischer, 1886), and Lycodon ruhstrati abditus (Vogel et al., 2009). In addition, the new species differs from L. cardamomensis by having more ventral scales (241 versus 215), and in body pattern (87 blotches versus 12 bands); from L. davidi by having more ventral scales (241 versus 224), six dorsal scale rows on the posterior third of the body feebly keeled (versus dorsal scale rows at midbody slightly keeled, outermost rows entirely smooth throughout body), and belly pattern uniform grey cream (versus anterior third whitishcream, posterior part heavily speckled with dark dots); from L. multifasciatus by having more ventral scales (241 versus maximum 237), and dorsal pattern blotched (versus banded); from L. ophiophagus by having more ventral scales (241 versus 211), and dorsal pattern (87 blotches versus 21-22 bands); from L. paucifasciatus by having fewer dorsal scale rows at neck (17 versus 19), more ventral scale rows (241 versus 221-222), six dorsal scale rows on the posterior third of the body feebly keeled (versus two upper rows plus vertebral row distinctly keeled), and dorsal pattern blotched (versus banded) (Neang et al., 2014); from L. r. ruhstrati and Lycodon ruhstrati abditus by having more ventral scales (241 versus 211-228; 241 versus 206-224, respectively), and dorsal pattern blotched (versus banded in the latter) (Vogel et al., 2012); from L. synaptor by having much more ventral scale rows (241 versus 201-203), dorsal pattern with 87 blotches (versus 30-31 bands), and belly pattern uniform grey cream (versus banded) (Vogel & David 2010); from L. zoosvictoriae by having more ventral scales (241 versus 213), dorsal pattern consisting of 87 blotches (versus 31), and having six dorsal scale rows on the posterior third of the body feebly keeled (versus all weakly keeled) (Neang et al., 2014). From the remaining species occurring in Laos, the new species can be distinguished as follows: from L. capucinus by having more ventrals (241 versus 182-211), fewer supralabials (8/8 versus 9-10), cloacal single (versus divided), dorsal blotches 87 (versus reticulated), and greyish yellow blotched body pattern (versus reticulated); from L. fasciatus by having more ventral scale rows (241 versus 182-225), dorsal pattern consisting of 87 blotches (versus 19-49 bands), and belly pattern uniform grey cream (versus white with dark blotches) (Neang et al., 2014); from L. laoensis by having loreal in contact with the orbit (versus separated), more ventrals (241 versus 169-192), and dorsal scales feebly keeled in the posterior body part (versus all smooth) (Neang et al., 2014); from L. septentrionalis by having more infralabials (10 versus 7-8), more ventral scales (241 versus 202-217), and dorsal pattern blotched (versus banded), as well as belly pattern uniform grey cream (versus white) (Neang et al., 2014); from L. subcinctus by the presence of preocular scale (versus absent), having cloacal scale single (versus divided), dorsal pattern blotched (versus banded in anterior part), and more ventral scale rows (241 versus 129-230) (Neang et al., 2014). From the remaining species in the fasciatus group, the new species differs as follows: from L. butleri Boulenger by having more ventral scale rows (241 versus 220-227), dorsal pattern blotched (versus banded), and belly pattern uniform grey cream (versus banded & spotted) (Grismer et al., 2014); from L. cavernicolus by having dorsal head uniformly dark grey (versus light brown), fewer supralabials (8 versus 9 or 10), more dorsal blotches (87 versus 36-45 bands), dorsal scales on the anterior 2/3 of the body length smooth, the six central dorsal scale rows of the posterior 1/3 of the body length feebly keeled (versus all keeled), and greyish yellow blotched pattern on the body (versus white bands); from L. gongshan by having six dorsal scale rows on the posterior third of the body feebly keeled (versus upper and vertebral dorsal rows keeled), more ventral scale rows (241 versus 210- 216), and dorsal pattern with 87 blotches (versus 32- 40 bands) (Vogel & Luo, 2011); from L. liuchengchaoi by having cloacal scale single (versus divided), dorsal pattern consisting of 87 irregular greyish yellow dorsal blotches (versus 40 well-defined yellow rings), and more ventral scales (241 versus 204) (Zhang et al., 2011). Distribution: Lycodon banksi sp. nov. is currently known only from the type locality in the Phou Hin Poun NPA, Khammouane Province, central Laos (Fig. 6). Etymology: The name of the species is dedicated to our friend and colleague Chris Banks, International Coordinator, Philippine Crocodile National Recovery Team, Zoos Victoria, Australia, for his outstanding contributions towards amphibian and reptile conservation, in particular of the Philippine Crocodile. We propose the following common names: Banks’ Wolf Snake (English), Banks Wolfszahnnatter (German). Natural history: The holotype was found at 20:39 h, crawling on a limestone outcrop in the karst forest, approximately 0.3 m above the forest floor, at an elevation of 167 m a.s.l. The humidity at the time of collection was approximately 85% and the air temperature ranged from 23 to 26 oC (Fig. 7). Another specimen was observed 12.4 km away from the type locality, active on the ground at 23:30 h, near a limestone cliff in the secondary forest.

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

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

Gekko nadenensis sp. nov. (Fig. 2) Holotype. VNUF R.2016.1, adult male, on a karst cliff, near the entrance of Nang Log cave (17��30���N, 105��23���E, elevation 178 m a.s.l.), Naden Village, Gnommalath District, Khammouane Province, central Laos, collected by Vinh Quang Luu and Kieusomphone Thanabuaosy on 20 February 2016. Paratype. NUOL-R.2016.2, adult male, the same data as the holotype; ZFMK 98741, gravid adult female, collected on a karst cliff inside Nang Log cave by Vinh Quang Luu, Thomas Calame, and Kieusomphone Thanabuaosy on 29 March 2015. Diagnosis. Gekko nadenensis sp. nov. differs from its relatives by a combination of the following characters: a medium-sized gecko species (SVL 61.0��� 77.1 mm); nares bordered with rostral; internasals absent; postmentals enlarged; interorbital scales between anterior corners of the eyes 28���30; dorsal tubercles absent; ventral scales between mental and cloacal slit 175���185; midbody scale rows 123���140; ventral scale rows 38���40; subdigital lamellae on first toes 13���15, on fourth toes 14���16; finger and toe webbing present at base; tubercles on upper surface of fore and hind limbs absent; precloacal pores six (3+3 or 5+1) in a discontinuous row in males and absent in the female; postcloacal tubercles 1 or 2; tubercles absent on dorsal surface of tail base; subcaudals distinctly enlarged; dorsal surface of body with greyish brown blotches. Description of holotype. An adult male with a total length of 134.1 mm (SVL 61.0 mm, TaL 73.1 mm); body slender, elongate (AG/SVL ratio 0.46); head longer than wide (HW/HL ratio 0.70); rostral quadrangular, nearly twice wider than high (RH/RW ratio 0.56) and wider than mental (MW/RW ratio 0.78 mm), bordered with first supralabial and supranasal on each side; nostrils round, touching rostral, first supralabial, supranasal, and two enlarged nasals posteriorly; posterior nasal region concave; internasal absent; preorbitals 22/22, preorbital region deeply concave; interorbitals 30; eye large (OD/HL ratio 0.24), pupil vertical; ear opening oval, oblique, smaller than eye (TD/OD ratio 0.44); mental triangular, wider than long (ML/MW ratio 0.86); enlarged postmentals two, hexagonal, twice as long as wide, and longer than length of mental, touching mental, first infralabial on both sides and seven gular scales posteriorly, outer gular scales larger than inner scales; supralabials 12/14; infralabials 10/10; dorsal scales on body smooth, round or oval, granular, juxtaposed; lateral fold distinct; ventrals distinctly larger than dorsal scales, smooth, imbricate, and largest in the middle of belly; ventral scales at midbody 40; scale rows around midbody 140; ventral scales in a row between mental and cloacal slit 175; scales on dorsal surface of forelimbs slightly enlarged; tubercles on dorsal surface of limbs absent; scales on anterior and ventral parts of femur larger than those on posterior and dorsal parts; enlarged femoral scales absent; fingers and toes basally webbed; subdigital lamellae under first fingers 15/15, under fourth finger 15/14, under first toe 13/14, under fourth toe 15/16; precloacal pores six (3+3), in a discontinuous row, precloacal scales enlarged; postcloacal tubercles 1/1; base of tail thickened, without tubercles on dorsal surface; subcaudals enlarged, smooth, imbricate. Coloration in life. Dorsal surface of head dark grey with grey blotches and brownish spots; posterior part of the orbit with a grey stripe; labials with grey and dark bars; crown region with a grey star aniseed shape; neck with three grey oval-shaped blotches; dorsal surface of body dark grey with grey blotches and brownish spots, largest blotches on the midbody; dorsal surface of fore and hind limbs brownish grey with light transverse bars; ventral surface of head, belly, and limbs cream with black dots; dorsal surface of tail with eight grey transverse bands, more distinct posteriorly; ventral tail grey-brown in forepart and with nearly closed bands in hindpart. Sexual dimorphism. Measurements and scalation of the female paratype are shown in Table 2. The female paratype has a larger size (SVL 77.1 mm versus 61.0��� 74.3 mm in males) and without precloacal pores. TABLE ��. Morphological comparisons among the species of the Gekko japonicus group (mođifieđ after Luu et al. 2015, abbreviations đefineđ in text, - = đata unavailable). ......continued on the next page TABLE ��. (continueđ) ......continued on the next page TABLE ��. (continueđ) Comparisons. Based on examination of specimens and data obtained from the literature (Boulenger 1907; Ota et al. 1995; R��sler et al. 2005, 2010, 2011; Yang et al. 2012, Nguyen et al. 2013; Luu et al. 2014; Ngo et al. 2015; Yang 2015; Luu et al. 2015) we compared the new species from Laos with the remaining members of the Gekko japonicus group (R��sler et al. 2011; Luu et al. 2015) (see Table 3). Morphologically, the new Gekko species can be distinguished from the species of the G. japonicus group (following R��sler et al. 2011; Luu et al. 2015) as follows: Gekko nadenensis sp. nov. does not have dorsal tubercles and thus differs from the following species which have tubercles on the back: G. adleri, G. auriverrucosus, G. canhi, G. chinensis, G. japonicus, G. hokouensis, G. kwangsiensis, G. liboensis, G. palmatus, G. scabridus, G. shibatai, G. similignum, G. swinhonis, G. vertebralis, and G. wenxianensis. The new species can be distinguished from G. aaronbaueri by having fewer interobital scales (28���30 versus 34���37), more scale rows around midbody (123���140 versus 98���104), and more precloacal pores in males (6 versus 3���4); from G. melli by its smaller size (SVL reaching 77.1 mm versus 84.6 mm), fewer interorbitals (28���30 versus 34���40), internasals absent (versus present), postmentals enlarged (versus not enlarged), and fewer precloacal pores in males (6 versus 9���11); from G. subpalmatus by having fewer interorbitals (28���30 versus 32), postmentals enlarged (versus not enlarged), the absence of internasals (versus present), having fewer ventral scales (38���40 versus 48), and precloacal pores in a discontinuous row in males (versus in a continuous row); from G. taibaiensis by its larger size (SVL reaching 77.1 mm versus 69 mm), having more supralabials (12���14 versus 9���10), and more lamellae under first and fourth toes (13���15 versus 6��7 and 14���16 versus 7 or 8, respectively); from G. tawaensis by the lack of internasals (versus 2), having postmentals enlarged (versus not enlarged in G. tawaensis), and precloacal pores present (versus absent); from G. truongi by its smaller size (SVL reaching 77.1 mm versus 95.9 mm), having fewer interobital scales (28���30 versus 45���48), more scales from mental to cloacal slit (175���185 versus 160���172), and fewer precloacal pores in males (6 versus 10���11); and from G. yakuensis by lacking internasals (versus having), having postmentals enlarged (versus not enlarged), fewer precloacal pores in males (6, in a discontinuous row versus 6���8, in a continuous row), and the absence of tubercles on dorsal surface of tail (versus presence). Gekko nadenensis sp. nov. is similar to G. bonkowskii, G. thakhekensis, G. scientiadventura, and G. sengchanthavongi in body size and dorsal pattern. However, the new species can be distinguished from G. bonkowskii by having more interorbitals (28���30 versus 26���27), precloacal pores in a discontinuous row in males (versus in a continuous row), more scales from mental to cloacal slit (175���185 versus 154���169), and more scale rows at midbody (123���140 versus 117); from G. thakhekensis by having more interorbitals (28���30 versus 22���26), more precloacal pores in males (6, in a discontinuous row versus 1���5, in a continuous row), more scales from mental to cloacal slit (175���185 versus 165���174), and more scale rows at midbody (123���140 versus 110���116); from G. scientiadventura by having fewer interorbitals (28���30 versus 41���51), and precloacal pores in a discontinuous row in males (versus in a continuous row), and more scales from mental to cloacal slit (175���185 versus 118���140); from G. sengchanthavongi by having more precloacal pores in males (6 versus 4���5), more supralabials and infralabials (12���14 versus 8��10 and 10���12 versus 6���7, respectively), dorsal head pattern with light grey blotches and a grey stripe behind the orbit (versus dark and grey spots), and dorsal body pattern with light grey blotches and brown flecks (versus irregular transverse dark brown bars). For more details see Table 4. Distribution. Gekko nadenensis sp. nov. is currently known only from the type locality in the karst forest of Nang Log cave, Naden Village, Gnommalath District, Khammouane Province, central Laos (Fig. 3). Etymology. We name this species after its type locality, Naden Village, to underscore the importance of the limestone forest in terms of biodiversity and nature conservation. From this site another new gecko species (Cyrtodactylus rufford) was described recently (Luu et al. 2016). We suggest as common names: Naden Gecko (English), Kap Ke Naden (Laotian), and Naden Gecko (German). Natural history. Specimens of the new species were collected between 19:30 and 22:00, on a karst cliff near the entrance of and inside Nang Log Cave, from 0.3 m to 3 m above the ground, at elevations between 158 and 178 m a.s.l. (Fig. 4). The humidity was approximately 70���80% and the air temperature ranged from 18 to 26o C. The new species co-occurs with the recently described bent-toed gecko Cyrtodactylus rufford (Luu et al. 2016). The female paratype, which was collected in March 2015 contained two eggs. In February 2016, we found a mass egglaying site (about 2 m above the ground) in a dark and narrow corner of a rock crevice system at the mouth of the Nang Log cave. Some of the eggs had not yet hatched (Fig. 5)., Published as part of Luu, Vinh Quang, Nguyen, Truong Quang, Le, Minh Duc, Bonkowski, Michael & Ziegler, Thomas, 2017, A new karst dwelling species of the Gekko japonicus group (Squamata: Gekkonidae) from central Laos, pp. 179-193 in Zootaxa 4263 (1) on pages 181-190, DOI: 10.11646/zootaxa.4263.1.10, http://zenodo.org/record/572614, {"references":["Luu, V. Q., Calame, T., Nguyen, T. Q., Le, M. D. & Ziegler, T. (2015) Morphological and molecular review of the Gekko diversity of Laos with descriptions of three new species. Zootaxa, 3986 (3), 279 - 306. https: // doi. org / 10.11646 / zootaxa. 3986.3.2","Boulenger, G. A. (1907) Description of new lizard in the British Museum. Annals and Magazine of Natural History, 19 (7), 486 - 489.","Ota, H., Lau, M. W., Weidenhofer, T., Yasukawa, Y. & Bogadek, A. (1995) Taxonomic review of the geckos allied to Gekko chinensis Gray 1842 (Gekkonidae, Reptilia) from China and Vietnam. Tropical Zoology, 8, 181 - 196. https: // doi. org / 10.1080 / 03946975.1995.10539278","Rosler, H., Ziegler, T., Vu, N. T., Herrmann, H. W. & Bohme, W. (2005 [\" 2004 \"]) A new lizard of the genus Gekko Laurenti, 1768 (Squamata: Sauria: Gekkonidae) from the Phong Nha-Ke Bang National Park, Quang Binh Province, Vietnam. Bonner zoologische Beitrage, 53 (1 / 2), 135 - 148.","Rosler, H., Nguyen, Q. T., Doan, V. K., Ho, T. C., Nguyen, T. T. & Ziegler, T. (2010) A new species of the genus Gekko Laurenti (Squamata: Sauria: Gekkonidae) from Vietnam with remarks on G. japonicus (Schlegel). Zootaxa, 2329, 56 - 68.","Rosler, H., Bauer, A. M., Heinicke, M., Greenbaum, E., Jackman, T., Nguyen, Q. T. & Ziegler, T. (2011) Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae). Zootaxa, 2989, 1 - 50.","Yang, J. - H., Wang, Y. - Y .., Zhang, T. - D., Sun, Y. - J. & Lin, S. - S. (2012) Genetic and morphological evidence on the species validity of Gekko melli Vogt, 1922 with notes on its diagnosis and range extension (Squamata: Gekkonidae). Zootaxa, 3505, 67 - 74.","Nguyen, Q. T., Wang, Y-Y. Yang, J-H., Lehmann, T., Le, M. D., Ziegler, T. & Bonkowski, M. (2013) A new species of the Gekko japonicus group (Squamata: Sauria: Gekkonidae) from the border region between China and Vietnam. Zootaxa, 3652 (5), 501 - 518.","Luu, V. Q., Calame, T., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2014) A new species of the Gekko japonicus group (Squamata: Gekkonidae) from central Laos. Zootaxa, 3895 (1), 73 ‾ 88.","Ngo, V. T., Pham, H. T., Phimvahan A., David, P. & Teynie, A. (2015) Gekko aaronbaueri, a new gecko (Squamata: Gekkonidae) from central Laos. Zootaxa, 3914 (2), 144 - 156.","Yang, J. - H. (2015) A new species of the genus Gekko Laurenti (Squamata: Sauria: Gekkonidae) from Guangxi, China. Zootaxa, 3936 (2), 287 - 295.","Luu, V. Q., Calame, T., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016) Cyrtodactylus rufford, a new cavedwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4067 (2), 185 - 199."]}

Cyrtodactylus sonlaensis sp. nov. (Figs. 3–5) Holotype. IEBR A.2017.1 (Field No. SL 2016.68), adult male, collected on 18 June 2016 by A.V. Pham and H.V. Tu, in the karst forest near Bang Village (21o05.700’N, 104o48.179’E, elevation 1050 m above sea level, asl.), Muong Bang Commune, Phu Yen District, Son La Province, northwestern Vietnam. Paratypes. TBU 2017.1 (Field No. SL2016.67, subadult male), IEBR A.2017.2 (Field No. SL 2016.69, adult female), the same data as the holotype; VNMN 2017.1 (Field No. SL 2016.467, adult female), collected on 28 October 2016 by A.V. Pham and T.Q.L. Hoang, in the karst forest near Bang Village (21o06.406’N, 104o47.810’E, elevation 890 m a.s.l.), Muong Bang Commune, Phu Yen District, Son La Province, northwestern Vietnam. Diagnosis. The new species can be distinguished from other members of the genus Cyrtodactylus by a combination of the following characters: medium size (SVL up to 83.2 mm); dorsal tubercles in 13–15 irregular rows; 34–42 ventral scale rows; ventrolateral folds present without interspersed tubercles; 15–17 enlarged femoral scales on each thigh; femoral pores 14 or 15 on each thigh of males, absent in females; precloacal pores 8, in a continuous row in males, absent in females; postcloacal tubercles 2 or 3; lamellae under toe IV 18–21; dorsal head with dark brown marking, oval and arched shape; nuchal loop discontinuous; five brown dorsal bands between limb insertions, third and fourth discontinuous; subcaudal scales transversely enlarged. Description of holotype. Adult male, snout-vent length (SVL) 77.5 mm; body elongate (AG/SVL 0.46); head distinguished from neck, elongate, depressed (HL/SVL 0.28, HW/HL 0.69, HH/HL 0.44); supranasals in contact with each other anteriorly, separated from each other by a small scale posteriorly; nares oval, surrounded by supranasal, rostral, first supralabial, and three postnasals; loreal region concave; snout long (SE/HL 0.41), round anteriorly, longer than diameter of orbit (OD/SE 0.67); snout scales small, round, granular, larger than those in frontal and parietal regions; eye large (OD/HL 0.28), pupils vertical; upper eyelid fringe with spinous scales; ear opening oval, obliquely directed, small in size (ED/HL 0.09); rostral wider than high (RH/RW 0.71) with a medial suture, bordered by first supralabial, nostril and supranasal on each side; mental triangular, as wide as rostral (RW 3.1 mm, MW 3.0 mm), wider than high (ML/MW 0.76); postmentals two, enlarged, in contact posteriorly, bordered by mental anteriorly, first infralabial laterally, and an enlarged chin scale posteriorly; supralabials 11/11; infralabials 11/9. Dorsal scales granular; dorsal tubercles round, 3 or 4 times larger than the size of adjoining scales, conical, present on occiput, back and tail base, each surrounded by 9–11 granular scales, in 14 or 15 irregular longitudinal rows at midbody; ventral scales smooth, medial scales 2 or 3 times larger than dorsal scales, round, subimbricate, largest posteriorly, in 34–36 longitudinal rows at midbody; lateral skin folds distinct, without tubercles; gular region with homogeneous smooth scales; ventral scales between mental and cloacal slit 189–193 (counted three times); precloacal groove absent; three rows of enlarged scales present in posterior region of pore-bearing scales; femoral pores bearing scales enlarged, in a continuous row with pore-bearing precloacal scales on the left side but separated from pore-bearing precloacal scales by 2 poreless femoral scales on the right side; femoral pores 15/14; precloacal pores eight, in a continuous row. Fore and hind limbs moderately slender (ForeaL/SVL 0.17, CrusL/SVL 0.24); dorsal surface of forelimbs covered by few slightly developed tubercles; dorsal surface of hind limbs covered by distinctly developed tubercles; interdigital webbing weakly developed; subdigital lamellae: finger I 14/15 (with 6/6 basally broadened lamellae), finger II 17/17 (7/7), finger III 16/18 (7/7), finger IV 18/17 (7/7), finger V 18/17 (8/7), toe I 15/16 (6/6), toe II 18/17 (7/7), toe III 17/18 (7/7), toe IV -/18 (8/7), toe V 20/19 (8/7). Tail complete, longer than snout-vent length (TaL 96.5 mm, Tal/SVL 1.24); postcloacal tubercles 3/3; dorsal tail base with distinct tubercles; subcaudals distinctly enlarged, smooth. Cyrtodactylus sonlaensis sp.nov. Cyrtodactylus otai......continued on the next page Cyrtodactylus sonlaensis sp.nov. Cyrtodactylus otai Coloration in preservative. Ground color of dorsal head and back greyish brown; snout region yellowish with three dark brown spots; dorsal head with dark brown marking, oval and arched shape; a dark stripe extending from posterior corner of eye rearwards to above tympanum; labials brown with cream sutures; neck with some large dark blotches, forming a discontinuous band anteriorly and a continuous band posteriorly; dorsum with five transverse dark brown bands between fore and hind limb insertions, edged in white anteriorly and posteriorly, third and fourth bands broken; dorsal surface of fore and hind limbs with dark brown blotches; tail greyish cream with ten dark brown bands; chin, throat, chest, belly and lower limbs cream; ventral surface of tail grey with seven dark brown bands. For coloration in life see Fig. 3. Sexual dimorphism and variation. The females differ from male specimens in the absence of precloacalfemoral pores and hemipenial swellings at the tail base. For other morphological characters see Table 2. Distribution. Cyrtodactylus sonlaensis sp. nov. is currently known only from the type locality in Phu Yen District, Son La Province, Vietnam (Fig. 1). Etymology. The specific epithet “ sonlaensis ” refers to the type locality, Son La Province, where the new species was discovered. Natural history. Specimens were found at night between 19:00 and 21:00, on trees near limestone cliffs and in rock crevices, about 0.1–2.0 m above the ground, at elevations between 900 and 1200 m a.s.l. The surrounding habitat was disturbed evergreen karst forest of medium hardwood and shrub. The humidity was approximately 80– 90% and the air temperature ranged from 22 to 29o C. Comparisons. We compared the new species with its congeners from Vietnam and neighboring countries in mainland Indochina, including Laos, Cambodia, Myanmar and Thailand based on examination of specimens (see Appendix) and data obtained from the literature (Smith 1917, 1921a, b, 1935; Taylor 1963; Ulber 1993; Bauer 2002, 2003; Bauer et al. 2002, 2003, 2009, 2010; Ziegler et al. 2002, 2010, 2013; Pauwels & Sumontha 2014; Pauwels et al. 2004, 2013, 2014a, b, 2016; Nguyen et al. 2006, 2014; Hoang et al. 2007; Orlov et al. 2007; Grismer & Ahmad; Ngo 2008; Ngo & Bauer 2008; Ngo & Grismer 2010, 2012; Ngo & Pauwels 2010; Ngo & Chan 2010, 2011; Ngo et al. 2008, 2010; Sumontha et al. 2010, 2012, 2014; Chan-ard & Makchai, 2011; David et al. 2011; Schneider et al. 2011; Luu et al. 2011, 2014, 2015, 2016a, b, c; Grismer et al. 2012; Kunya et al. 2014, 2015; Nazarov et al. 2014; Panitvong et al. 2014; Le et al. 2016; and Connette et al. 2017). Cyrtodactylus sonlaensis sp. nov. has distinctly enlarged subcaudals, which are only slightly or not enlarged in the following species: C. ayeyarwadyensis Bauer, C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. bobrovi Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, C. brevidactylus Bauer, C. brevipalmatus (Smith), C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. buchardi David, Teynié & Ohler, C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler, C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler, C. cucdongensis Schneider, Phung, Le, Nguyen & Ziegler, C. dati Ngo, C. gunungsenyumensis Grismer, Wood, Anuar, Davis, Cobos & Murdoch, C. gansi Bauer, C. hitchi Riyanto, Kurniati & Engilis, C. huynhi Ngo & Bauer, C. irregularis (Smith), C. mandalayensis Mahony, C. martini Ngo, C. otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, C. papilionoides Ulber & Grossmann, C. phuocbinhensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, C. pseudoquadrivirgatus Rösler, Vu, Nguyen, Ngo & Ziegler, C. quadrivirgatus Taylor, C. ranongensis Sumontha, Pauwels, Panitvong, Kunya & Grismer, C. slowinskii Bauer, C. sommerladi Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler, C. tamaiensis (Smith), C. taynguyenensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, C. thuongae Phung, Van Schingen, Ziegler & Nguyen, C. vilaphongi Schneider, Nguyen, Duc Le, Nophaseud, Bonkowski & Ziegler, C. wakeorum Bauer, C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler, C. ziegleri Nazarov, Orlov, Nguyen & Ho. Cyrtodactylus sonlaensis sp. nov. differs from C. aequalis Bauer by the absence of precloacal pores in females (vs. 9), fewer dorsal tubercle rows (13–15 vs. 24), and more ventral scale rows (34–42 vs. 24); from C. annandalei Bauer by its larger size (SVL 71.2–83.2 mm vs. 49.0–55.0 mm), more precloacal-femoral pores in males (37 vs. 12–24), and fewer dorsal tubercle rows (13–15 vs. 16–18); from C. angularis (Smith) by having more precloacal pores in males (8 vs. 3), the absence of precloacal pores in females (vs. 3), and the presence of femoral pores in males (vs. absence); from C. astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels by having fewer dark caudal bands (10 vs. 13–14); from C. auribalteatus Sumontha, Panitvong & Deein by having more enlarged femoral scales on each thigh (15–17 vs. 5– 7), more femoral pores on each thigh in males (14–15 vs. 4–5), and more precloacal pores in males (8 vs. 6); from C. badenensis Nguyen, Orlov & Darevsky by having more ventral scale rows (34–42 vs. 25–29), the presence of enlarged femoral scales (vs. absence), and the presence of precloacal-femoral pores in males (vs. absence); from C. bichnganae Ngo & Grismer by its smaller size (SVL 71.2–83.2 mm vs. 95.3–99.9 mm), having more enlarged femoral scales on each thigh (15–17 vs. 11–13), more femoral pores on each thigh in males (14–15 vs. 9), and fewer precloacal pores in males (8 vs. 10); from C. bansocensis Luu, Nguyen, Le, Bonkowski & Ziegler by having more precloacal-femoral pores in males (37 vs. 34), more infralabials (9–11 vs. 8), and more ventral scales at midbody (189–202 vs. 158–170); from C. calamei Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler by having fewer postcloacal tubercles (2–3 vs. 4) and the absence of precloacal-femoral pores in females (vs. 38); from C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen by its smaller size (SVL 71.2–83.2 mm vs. 90.4–94.0 mm), having more enlarged femoral scales on each thigh (15–17 vs. 8), more femoral pores on each thigh in males (14–15 vs. 6), fewer lamellae under finger IV (17–19 vs. 22) and under toe IV (18–21 vs. 23–25); from C. chanhomeae Bauer, Sumontha & Pauwels by having more precloacal-femoral pores in males (37 vs. 32) and the absence of precloacal-femoral pores in females (vs. 34); from C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau by its smaller size (SVL 71.2–83.2 mm vs. 90.9–99.3 mm), the presence of enlarged femoral scales (vs. absence), the presence of femoral pores in males (vs. absence), having more precloacal pores in males (8 vs. 6), and the absence of precloacal pores in females (vs. 7); from C. chrysopylos Bauer by having fewer precloacal pores in males (8 vs. 10) and the presence of femoral pores in males (vs. absence); from C. condorensis (Smith) by having more precloacal pores in males (8 vs. 4–7) and a different dorsal color pattern (banded vs. blotched); from C. consobrinoides (Annandale) by its larger size (SVL 71.2–83.2 mm vs. 48.0 mm), more precloacal pores in males (8 vs. 4), and more ventral scale rows (34–42 vs. 24–30); from C. cucphuongensis Ngo & Chan, by its smaller size (SVL 71.2–83.2 mm vs. 96.0 mm), the presence of precloacal-femoral pores in males (vs. absence), having fewer lamellae under finger IV (17–19 vs. 21), and under toe IV (18–21 vs. 24); from C. darevskii Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov by having fewer precloacalfemoral pores in males (37 vs. 38–44) and the absence of precloacal-femoral pores in females (vs. 24–34); from C. doisuthep Kunya, Panmongkol, Pauwels, Sumontha, Meewasana, Bunkhwamdi & Dangsri by its smaller size (SVL reaching 83.2 mm vs. 90.5 mm), having more precloacal pores in males (8 vs. 6) and fewer dorsal tubercle rows (13–15 vs. 19–20); from C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya by having more precloacal-femoral pores in males (37 vs. 17–19), the absence of precloacal-femoral pores in females (vs. 0–7), and more lamellae under finger IV (17–19 vs. 16); from C. eisenmanae Ngo by having fewer ventral scale rows (34–42 vs. 44–45), more enlarged femoral scales on each thigh (15–17 vs. 4–6), and the presence of precloacal-femoral pores in males (vs. absence); from C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya by having more ventral scale rows (34–42 vs. 28), fewer precloacal pores in males (8 vs. 9), more lamellae under finger IV (17–19 vs. 16), and a different dorsal color pattern (banded vs. blotched); from C. feae Boulenger by its larger size (SVL 71.2–83.2 mm vs. 45.0 mm) and the presence of precloacal-femoral pores in males (vs. absence); from C. grismeri Ngo by the presence of precloacal-femoral pores in males (vs. absence) and the presence of enlarged femoral scales (vs. absence); from C. hinnamnoensis Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler by its smaller size (SVL reaching 83.2 mm vs. 100.6 mm) and having fewer postcloacal tubercles (2–3 vs. 4–5); from C. hontreensis Ngo, Grismer & Grismer by having more enlarged femoral scales on each thigh (15–17 vs. 2–5); from C. huongsonensis Luu, Nguyen, Do & Ziegler by its smaller size (SVL reaching 83.2 mm vs. 89.8 mm), having more enlarged femoral scales on each thigh (15–17 vs. 7–9), more precloacal-femoral pores in males (37 vs. 23), and the absence of precloacal-femoral pores in females (vs. 23); from C. interdigitalis Ulber by having fewer femoral pores on each thigh in males (14–15 vs. 16–18) and fewer precloacal pores in males (8 vs. 14); from C. intermedius (Smith) by having more enlarged femoral scales on each thigh (15–17 vs. 6–10), the presence of femoral pores (vs. absent), having fewer lamellae under finger IV (17– 19 vs. 20) and under toe IV (18–21 vs. 22); from C. inthanon Kunya, Sumontha, Panitvong, Dongkumfu, Sirisamphan & Pauwels by having fewer dorsal tubercle rows (13–15 vs. 18–20), more femoral pores on each thigh in males (14–15 vs. 6), and more precloacal pores in males (8 vs. 5); from C. jaegeri Luu, Calame, Bonkowski, Nguyen & Ziegler by having more ventral scale rows (34–42 vs. 31–32), fewer precloacal-femoral pores in males (37 vs. 44), and the absence of precloacal-femoral pores in females (vs. 24); from C. jarujini Ulber, by its smaller size (SVL 71.2–83.2 mm vs. 85.0–90.0 mm), having fewer precloacal-femoral pores in males (37 vs. 52–54), generally more lamellae under finger IV (17–19 vs. 15–17), and a different dorsal color pattern (banded vs. blotched); from C. khammouanensis Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov by having fewer precloacal-femoral pores in males (37 vs. 40–44); from C. khelangensis Pauwels, Sumontha, Panitvong & Varaguttanonda by having more precloacal pores in males (8 vs. 2–5), fewer dorsal tubercle rows (13–15 vs. 16–20), more femoral pores on each thigh in males (14–15 vs. 6) and the absence of precloacal pores in females (vs. 6); from C. kingsadai Ziegler, Phung, Le & Nguyen by having more enlarged femoral scales on each thigh (15–17 vs. 9–12), more femoral pores on each thigh in the males (14–15 vs. 1–4), and the absence of precloacal pores in females (vs. 4–8); from C. kunyai Pauwels, Sumontha, Keeratikiat & Phanamphon by having more precloacal pores in males (8 vs. 3) and fewer dorsal tubercle rows (13–15 vs. 16–20); from C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels by the absence of precloacal-femoral pores in females (vs. 33–43); from C. lenya Mulcahy, Thura & Zug by the presence of precloacal-femoral pores in males (vs. absence) and more ventral scale rows (34–42 vs. 29); from C. lomyenensis Ngo & Pauwels by having fewer precloacal-femoral pores in males (37 vs. 39–40) and the absence of precloacal-femoral pores in females (vs. 32); from C. macrotuberculatus Grismer & Ahmad by its smaller size (SVL reaching 83.2 mm vs. 120.0 mm), more ventral scale rows (34–42 vs. 17–28) and fewer dorsal tubercle rows (13–15 vs. 19–27); from C. multiporus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov by having fewer precloacal-femoral pores in males (37 vs. 58–60); from C. nigriocularis Nguyen, Orlov & Darevsky by having more precloacal pores in males (8 vs. 0–2), the presence of enlarged femoral scales (vs. absence), the presence of femoral pores in males (vs. absence), and a different dorsal color pattern (banded vs. uniformly brown); from C. oldhami (Theobald) by having more precloacal pores in males (8 vs. 1–4), the presence of femoral pores in males (vs. absence), and a different dorsal color pattern (banded vs. striped and spotted); from C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler by having more precloacal pores in males (8 vs. 4), the presence of enlarged femoral scales (vs. absence), the presence of femoral pores in males (vs. absence), and the absence of precloacal pores in females (vs. 4); from C. payarhtanensis Mulcahy, Thura & Zug by the presence of precloacal-femoral pores in males (vs. absence) and having more ventral scale rows (34–42 vs. 26– 32); from C. peguensis Boulenger by the presence of femoral pores in males (vs. absence) and a diffe

Luu, Vinh Quang, Calame, Thomas, Nguyen, Truong Quang, Le, Minh Duc, Bonkowski, Michael, Ziegler, Thomas (2016): Cyrtodactylus rufford, a new cave-dwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa 4067 (2): 185-199, DOI: http://doi.org/10.11646/zootaxa.4067.2.4

Cyrtodactylus rufford sp. nov. (Figs. 2���3) Holotype. VFU R. 2015.14, adult male, on a karst cliff, near the entrance of Nang Log cave (17 �� 30.282 ���N, 105 �� 23.107 ���E, elevation 160 m a.s.l.), Gnommalath District, Khammouane Province, central Laos, was collected on 29 March 2015 by V. Q. Luu, T. Calame and K. Thanabuaosy. Paratypes. IEBR R. 2015.35, adult male, and NUOL R. 2015.15, subadult male; same collection data as was provided for the holotype. Diagnosis. The new species differs from other congeners of the genus Cyrtodactylus by the following combination of characters: medium size, SVL reaching 72.5 mm; dorsal pattern consisting of three or four light transverse bands between limb insertions; one intersupranasal; 14���16 irregular dorsal tubercle rows at midbody, weakly developed in the paravertebral region; 27���29 ventral scale rows between ventrolateral folds; 42���43 precloacal and femoral pores in a continuous row in males, enlarged femoral and precloacal scales present; 4 or 5 postcloacal tubercles on each side; dorsal tubercles present at base of tail; subcaudal scales medially enlarged. Description of the holotype. Adult male, SVL 68.3 mm; body slender, elongate (TrunkL/SVL ratio 0.47); head elongate (HL/SVL ratio 0.29), relatively narrow (HW/HL ratio 0.57), depressed (HH/HL ratio 0.36), distinct from neck; loreal region inflated, posterior nasal region concave; snout long (SE/HL ratio 0.47), obtuse, longer than orbit diameter (OD/SE ratio 0.57); scales on snout small, rounded, homogeneous, granular, larger than those on frontal and parietal regions; eye large (OD/HL ratio 0.27), pupils vertical; supraciliaries with tiny spines posteriorly; ear opening oval, small (EarL/HL ratio 0.10); rostral wider than high with a shallow inverted ��-shaped medial suture; supranasals separated from each other by one small scale; rostral in contact with first supralabial and nostril on each side; nostrils oval, each surrounded by supranasal, rostral, first supralabial, and two enlarged postnasals; mental triangular, as wide as rostral (RW 2.8, MW 2.7); one pair of enlarged postmentals, in broad contact posteriorly, bordered laterally by first infralabials and triangular mental anteriorly; supralabials 11 / 12; infralabials 10 / 11. Dorsal scales granular to weakly conical; intermixed with conical, round dorsal tubercles (2���3 times larger than adjoining scales) present from occipital region to dorsum and base of tail, tubercles in 16 rows at midbody, larger on flanks, each surrounded by 9 granular scales; ventrals larger than lateral scales, smooth, round, largest posteriorly, in 28 longitudinal scale rows between ventrolateral folds; gular region with homogeneous, smooth scales; ventral scales from mental to cloacal slit 153; precloacal groove lacking; enlarged femoral and cloacal scales present; femoral and precloacal pores 43, in a continuous row. Fore and hind limbs moderately slender (ForeL/SVL ratio 0.17, CrusL/SVL 0.20); dorsum of forelimbs with weakly developed tubercles; hind limbs dorsally with distinct tubercles; interdigital webbing slightly developed; lamellae under fourth finger 19 / 19; lamellae under fourth toe 19 / 18. Tail partly regenerated, longer than SVL (TaL 94.5 mm, TaL/SVL ratio 1.38); 5 / 4 postcloacal tubercles; tubercles present at base of tail dorsum; subcaudals transversely enlarged in the middle, flat, smooth. Coloration in life. Head dorsally greyish brown with dark blotches; nuchal loop brown, uninterrupted, in U��� shape; labials grey; three light transverse bands between limb insertions, edged in black anteriorly, somewhat irregular in the posterior parts of the dorsal bands; tubercles at midbody brown; tubercles on body bands grey; dorsal surface of fore- and hind- limbs with dark reticulated markings; dorsal surface of tail brown with light rings (2���3 times narrower than dark bands), ventral surface of head, body and limbs greyish cream; venter of tail grey. Variation: The adult male paratype (IEBR R. 2015.35) has wider light bands on the midbody with dark spots within each band compared to those of the holotype and subadult male paratype (NUOL R. 2015.15). Further morphological characters of the paratypes are provided in Table 3. Comparisons. We compared Cyrtodactylus rufford sp. nov. to other species of Cyrtodactylus from Laos and neighbouring countries in the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on examination of specimens (see Appendix) and data provided from taxonomic publications (Luu et al. 2014; Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014; Sumontha et al. 2015; Nguyen et al. 2015; Luu et al. 2015) (see Table 4). Cyrtodactylus rufford sp. nov. has distinctly enlarged median subcaudals and thus differs from the following species which are lacking enlarged median subcaudals: C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. bobrovi Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, C. buchardi David, Teyni�� & Ohler, C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. cattienensis Geissler, Nazarov, Orlov, B��hme, Phung, Nguyen & Ziegler, C. cucdongensis Schneider, Phung, Le, Nguyen & Ziegler, C. cryptus Heidrich, R��sler, Vu, B��hme & Ziegler, C. huynhi Ngo & Bauer, C. irregularis (Smith), C. otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, C. phuocbinhensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, C. pseudoquadrivirgatus R��sler, Vu, Nguyen, Ngo & Ziegler, C. quadrivirgatus Taylor, C. ranongensis Sumontha, Pauwels, Panitvong, Kunya & Grismer, C. taynguyenensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, C. thuongae Phung, van Schingen, Ziegler & Nguyen, C. vilaphongi Schneider, Nguyen, Le, Nophaseud, Bonkowski & Ziegler, and C. ziegleri Nazarov, Orlov, Nguyen & Ho. Cyrtodactylus rufford sp. nov. has femoral and precloacal pores in males, both of which are lacking in the following species: C. angularis (Smith), C. badenensis Nguyen, Orlov & Darevsky, C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, C. cucphuongensis Ngo & Chan, C. eisenmanae Ngo, C. grismeri Ngo, C. martini Ngo, C. nigriocularis Nguyen, Orlov & Darevsky, C. oldhami (Theobald), C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler, C. paradoxus (Darevsky & Szczerbak), C. puhuensis Nguyen, Yang, Le, Nguyen, Orlov, Hoang, Nguyen, Jin, Rao, Hoang, Che, Murphy & Zhang, C. saiyok Panitvong, Sumontha, Tunprasert & Pauwels, C. samroiyot Pauwels & Sumontha, C. sanook Pauwels, Sumontha, Latinne & Grismer, TABLE 4. Morphological comparisons between Cyrtodactylus rufford sp. nov. and its congeners from Laos and neighbouring countries in the Indochina region (compiled after Luu et al. 2014; Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014; Sumontha et al. 2015; Nguyen et al. 2015; Luu et al. 2015). Abbreviations are as follows: FPl = femoral pores in the left side; FPr = femoral pores in the right side; ��� = characters unobtainable from literature; * = tail regenerated, for other abbrevi- ations see material and methods. . darevskii 84.6 ���100.0 95.0���113.0 38���46 present present 38���44 24���34 17���20 18���22 banded present (FP+PP) . dumnuii 76.2���84.2 100.2 * 40 present present in 6 + 5���6 + 6���7 0��� 7 16 19 banded present males/abse (FPl+PP+FPr) nt in females ������continued on the next page TABLE 4. (Continued) TABLE 4. (Continued) C. spelaeus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov, C. sumonthai Bauer, Pauwels & Chanhome, C. wayakonei Nguyen, Kingsada, R��sler, Auer & Ziegler, C. takouensis Ngo & Bauer, C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome, and C. yangbayensis Ngo & Chan. Cyrtodactylus rufford sp. nov. has a contiguous series of femoral and precloacal pores in males and thus differs from the following species which have poreless scales separating femoral from precloacal pores: C. bichnganae Ngo & Grismer, C. brevipalmatus (Smith), C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya, C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya, C. huongsonensis Luu, Nguyen, Do & Ziegler, C. interdigitalis, C. khelangensis Pauwels, Sumontha, Panitvong & Varaguttanonda, and C. tigroides Bauer, Sumontha & Pauwels. Cyrtodactylus rufford sp. nov. has 42���43 femoral and precloacal pores in males and thus differs from the following species which have distinctly fewer femoral and precloacal pores: C. auribalteatus Sumontha, Panitvong & Deein (10���11), C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen (15), C. chanhomeae Bauer, Sumontha & Pauwels, C. intermedius (Smith) (8���10), C. kingsadai Ziegler, Phung, Le & Nguyen (7���16), C. roesleri (20���28), and C. soudthichaki (29). Cyrtodactylus rufford sp. nov. can be distinguished from C. astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels by its smaller size (maximum SVL 72.5 mm versus 108.3 mm), having fewer ventral scale rows (27���29 versus 31���46), fewer lamellae under the fourth toe (18��� 19 versus 20���24), and more femoral and precloacal pores in males (42���43 versus 31���38); from C. jarujini Ulber by its smaller size (maximum SVL 72.5 mm versus 90.0 mm), having fewer ventral scale rows (27���29 versus 32���38), and fewer femoral and precloacal pores in males (42���43 versus 52���54); from C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels by its smaller size (SVL reaching 72.5 mm versus 103.5 mm), having fewer ventral scales (27���29 versus 31���43), and more femoral and precloacal pores in males (42���43 versus 30���36); from C. multiporus by its smaller size (maximum SVL 72.5 mm versus 98.0 mm), having fewer ventral scale rows (27���29 versus 30���38), and fewer femoral and precloacal pores in males (42��� 43 versus 58���60); from C. teyniei David, Nguyen, Schneider & Ziegler by its smaller size (maximum SVL 72.5 mm versus 89.9 mm), having fewer ventral scale rows (27���29 versus 38), the presence of a nuchal band (versus being absent), and having a banded dorsal pattern (versus blotched). Cyrtodactylus rufford sp. nov. is similar to the members of the C. phongnhakebangensis group (Nazarov et al. 2014) including C. khammouanensis, C. lomyenensis, C. jaegeri, C. darevskii, and C. phongnhakebangensis Ziegler, R��sler, Herrmann & Vu in having a high number of femoral and precloacal pores and in the dorsal pattern. However, the new species can be distinguished from C. khammouanensis by having fewer ventral scale rows (27��� 29 versus 32���38), fewer dorsal tubercles at midbody (14���16 versus 16���21), fewer lamellae under the fourth toe (18���19 versus 20���23), dorsum of head with dark blotched markings (versus absent), having light transverse bands between limb insertions with wavy margins (versus transverse bands with smooth margins), and tail with light rings (versus light bands); from C. lomyenensis by having fewer ventral scale rows (27���29 versus 35���36), fewer supralabials (10���12 versus 13���14), and more femoral and precloacal pores in males (42���43 versus 39���40); from C. darevskii by its smaller size (maximum SVL 72.5 mm versus 100.0 mm), having fewer ventral scale rows (27���29 versus 38���46), and fewer ventral scales between mental and cloacal slit (153���167 versus 180���208); from C. jaegeri by having fewer ventral scale rows (27���29 versus 31���32), fewer femoral and precloacal pores in males (42���43 versus 44), fewer lamellae under the fourth toe (18���19 versus 20���23), and having irregular shaped light transverse bands between limb insertions (versus regular shaped light transverse bands); and from C. phongnhakebangensis by its smaller size (maximum SVL 72.5 mm versus 96.3 mm), having fewer ventral scale rows (27���29 versus 32��� 42), generally more femoral and precloacal pores in males (42���43 versus 32���42), and fewer ventral scales between mental and cloacal slit (153���167 versus 180���208). For more details see Table 5. Distribution. Cyrtodactylus rufford sp. nov. is currently known only from the type locality in the karst forest near Daen Village, Gnommalath District, Khammouane Province, central Laos (Fig. 4). Etymology. The new species is named in honour of the Rufford Foundation (UK) for its support to herpetofaunal research and conservation in Laos. The species epithet is to be treated as a noun in apposition, invariable. As common names, we suggest Ki Chiem Rufford (Laotian) and Rufford Bent-toed Gecko (English). Natural history. The type series of the new species was found between 20:00 and 21:00, on a karst cliff near the entrance of Nang Log Cave, from 0.3 m to 2 m height above the forest floor, at elevations between 160 and 180 m a.s.l. Nang Log Cave is situated within an isolated karst mountain surrounded by secondary forest, plantations, and some huts for tourists. The cave is a tourism site, only 50 m distant from Road 12 and 64 km from the border of Vietnam. The humidity was approximately 80 % and the air temperature ranged from 24 to 26 o C (Fig. 5). Other species found on the same karst wall within a 300 meters range of the type locality were the lizard Gekko gecko (Linnaeus), and the tree frog species Rhacophorus spelaeus Orlov, Gnophanxay, Phimminith & Phomphoumy, as well as the Huntsman Spider Heteropoda maxima Jaeger, and the long-legged cave centipede Thereuopoda longicornis (Fabricius). Transverse dorsal bands 4 light bands 3-4 sometimes irregular shaped 2���3 light bands between limbs light bands Tail pattern light rings light bands light bands, Published as part of Luu, Vinh Quang, Calame, Thomas, Nguyen, Truong Quang, Le, Minh Duc, Bonkowski, Michael & Ziegler, Thomas, 2016, Cyrtodactylus rufford, a new cave-dwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos, pp. 185-199 in Zootaxa 4067 (2) on pages 188-196, DOI: 10.11646/zootaxa.4067.2.4, http://zenodo.org/record/265253, {"references":["Luu, V. Q., Calame, T., Bonkowski, M., Nguyen, T. Q. & Ziegler, T. (2014) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Khammouane Province, Laos. Zootaxa, 3760 (1), 54 - 66. http: // dx. doi. org / 10.11646 / zootaxa. 3760.1.3","Nazarov, R., Poyarkov, N. A., Orlov, N. I., Nguyen, S. N., Milto, K. D., Martynov, A. A., Konstantinov, E. L. & Chulisov, A. S. (2014) A review of genus Cyrtodactylus (Reptilia: Sauria: Gekkonidae) in fauna of Laos with description of four new species. Proceedings of the Zoological Institute RAS, 318 (4), 391 - 423.","Nguyen, S. N., Yang, J. - X., Le, T. - N. T., Nguyen, L. T., Orlov, N. L., Hoang, C. V., Nguyen, T. Q., Jin, J. - Q., Rao, D. - Q., Hoang, T. N., Che, J., Murphy, R. W. & Zhang, Y. - P. (2014) DNA barcoding of Vietnamese bent-toed geckos (Squamata: Gekkonidae: Cyrtodactylus) and the description of a new species. Zootaxa, 3784 (1), 48 - 66. http: // dx. doi. org / 10.11646 / zootaxa. 3784.1.2","Schneider, N., Nguyen, T. Q., Le, M. D, Nophaseud, L., Bonkowski, M. & Ziegler, T. (2014) A new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of northern Laos. Zootaxa, 3835 (1), 080 - 096. http: // dx. doi. org / 10.11646 / zootaxa. 3835.1.4","Sumontha, M., Pauwels, O. S. G., Panitvong, N., Kunya, K. & Grismer, J. L. (2015) A new lowland forest Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Ranong Province, peninsular Thailand. Zootaxa, 3911 (1), 106 - 118. http: // dx. doi. org / 10.11646 / zootaxa. 3911.1.6","Luu, V. Q., Calame, T., Nguyen, T. Q., Bonkowski, M. & Ziegler, T. (2015) A new species of Cyrtodactylus (Squamata: Gekkonidae) from limestone forest, Khammouane Province, central Laos. Zootaxa, 4058 (3), 388 - 402. http: // dx. doi. org / 10.11646 / zootaxa. 4058.3.6"]}

Cyrtodactylus hinnamnoensis sp. nov. (Fig. 8) Cyrtodactylus phongnhakebangensis— Luu, Nguyen, Calame, Hoang, Soudthichack, Bonkowski, Ziegler, 2013. Biodiversity Data Journal, e 1015: 4. Holotype. IEBR A. 2013.90, adult male, from Ban Dou Village (17 ° 30.385 ’N, 105 ° 49.160 ’E, elevation 183 m a.s.l.) within Hin Nam No NPA, Khammouane Province, central Laos, was collected on 11 June 2013 by V. Q. Luu and N. V. Ha. Paratypes. IEBR A. 2013.89, adult male, 7 May 2013, from Hang Toi region, Noong Ma Village (17 ° 17.766 ’N, 106 °08.803’E, elevation 580 m a.s.l.); VNUF R. 2013.1 and NUOL R- 2013.2, adult males, 9 June 2013, from Vang Ma No Village (17 ° 30.778 ’N, 105 ° 49.259 ’E, elevation 180 m a.s.l.); VNUF R. 2014.99, adult male, 27 May 2014, from Cha Lou Village (17 ° 19.504 ’N, 105 ° 57.630 ’E, elevation ca. 300 m a.s.l.); ZFMK 95235, adult female, 8 May 2013, from Hang Toi region, Noong Ma Village (17 ° 17.763 ’N, 106 °08.778’E, elevation 555 m a.s.l.); ZFMK 95236, adult female, 30 May 2013, from Noong Choong Region, Cha Lou Village (17 ° 20.248 ’N, 105 ° 56.693 ’E, elevation 252 m a.s.l.); NUOL R- 2013.3, adult female, 11 June 2013, from Ban Dou Village (17 ° 31.545 ’N, 105 ° 49.086 ’E, elevation 197 m a.s.l.); VNUF R. 2015.3, female, 13 March 2015, from Xebangfai cave, Noong Ping Village (17 ° 22.459 ’N, 105 ° 49.626 ’E, elevation 182 m a.s.l.); NUOL R- 2015.9, female, 13 March 2015, from Xebangfai cave, Noong Ping Village (17 ° 22.648 ’N, 105 ° 52.931 ’E, elevation 182 m a.s.l.); VNUF R. 2015.11, female, 14 March 2015, from Xebangfai cave, Noong Ping Village (17 ° 22.759 ’N, 105 ° 52.931 ’E, elevation 285 m a.s.l.). The paratypes (VNUF R. 2015.3, NUOL R- 2015.9, and VNUF R. 2015.11) were collected by V. Q. Luu and K. Thanabuaosy in March 2015; the paratype (VNUF R. 2014.99) was collected by V. Q. Luu, N. V. Ha, T. Calame, D. V. Phan and K. Thanabuaosy in May 2014, the remaining type series was collected by V. Q. Luu, N. V. Ha, and K. Thanabuaosy in May and June 2013 (V. Q. Luu et al.). Diagnosis. Cyrtodactylus hinnamnoensis sp. nov. is characterized by: Adult SVL 84.1 ± 11.7 mm (mean ± SD); dorsal head with dark blotches; nuchal loop wide, distinct, posteriorly enlarged; dorsal body with four to six blackish brown bands between limb insertions; 13–19 irregular, weakly keeled dorsal tubercle rows; 35–48 ventral scale rows; ventral scale rows from mental to cloacal slit 179–201; scale rows at midbody 93–112; ventrolateral folds present, without tubercles; 36–44 precloacal-femoral pores in the males; 0–28 pores in females; enlarged femoral and precloacal scales present; 4–6 postcloacal tubercles; subcaudals enlarged. Description of holotype. Adult male, snout-vent length (SVL) 83.6 mm; body elongate (TrunkL/SVL 0.42); head elongate (HL/SVL 0.27), relatively wide (HW/HL 0.68), depressed (HH/HL 0.40), distinguished from neck; loreal region concave; snout long (SE/HL 0.43), obtuse, longer than diameter of orbit (OD/SE 0.60); snout scales small, homogeneous, granular, larger than those on frontal and parietal regions; eye large (OD/HL 0.26), pupils vertical; eyelid fringe with tiny spines posteriorly; ear oval-shaped, small (EarL/HL 0.08); rostral wider than high (RH/RW 0.60), with a median suture; supranasals in contact anteriorly and separated from each other by a small scale posteriorly; rostral in contact with first supralabial and nostril scales on each side; nares oval, surrounded by supranasal, rostral, first supralabial, and two enlarged postnasals; mental triangular, wider than high (ML/MW 0.65); two enlarged postmentals in broad contact posteriorly, bordered by mental anteriorly, first two infralabials laterally, and eight small scales posteriorly; supralabials 12 / 10; infralabials 10 / 9. Dorsal scales granular to flattened; dorsal tubercles round, weakly keeled, present on occiput, back and tail base, each surrounded by 8 granular scales, in 15 irregular longitudinal rows at midbody; ventral scales smooth, medial scales 2–3 times larger than dorsal scales, round, subimbricate, largest posteriorly, in 35 longitudinal rows between lateral folds at midbody; ventrolateral folds present, without tubercles; gular region with homogeneous smooth scales; ventral scales in a line from mental to cloacal slit 186; precloacal groove absent; enlarged femoral scales present; femoral and precloacal pores 42. Fore and hind limbs moderately slender (ForeL/SVL 0.17, CrusL/SVL 0.21); dorsal fore limbs with slightly developed tubercles; dorsal hind limbs covered by distinctly developed tubercles; fingers and toes free of webbing; lamellae under fourth fingers 19 / 19, under fourth toes 20 / 20. Tail regenerated, postcloacal tubercles 5 / 5; dorsal tail bearing tubercles at base; subcaudals distinctly enlarged, flat, smooth. Coloration in life. Ground coloration of dorsal head greyish brown with dark blotches; nuchal loop black, in U-shape, from posterior corner of eye through tympanum to the neck, dark brown, edged in yellow; body bands between limb insertions five, somewhat irregular, dark brown, edged in white; dorsal surface of fore and hind limbs with dark bars; tail brown dorsally with light brown rings, edged by yellowish white; chin, throat, and belly cream; upper and lower lips with dark brown bars; tail ventrally grey with light dots. Sexual dimorphism. The females differ from the males by lacking or having fewer precloacal-femoral pores (0–28 versus 36–44 in the males) and the absence of hemipenial swellings at the tail base (see Table 6). Comparisons. We compared the new species with its congeners from Laos and neighbouring countries in the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on examination of specimens (see Appendix) and data integrated from the literature (compiled after Luu et al. 2014 a; Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014; Sumontha et al. 2015; Nguyen et al. 2015; Luu et al. 2015 a; Luu et al. 2016 a, b) (see Tables 3). The cluster and correspondence analyses of morphological characters supported Cyrtodactylus hinnamnoensis sp. nov. as a sister taxon to C. darevskii (Figs. 2–3). Molecular phylogenetic analyses also demonstrated the close relationships between these species (see Fig. 1). Morphologically, Cyrtodactylus hinnamnoensis sp. nov. is closely related to the C. phongnhakebangensis group including C. darevskii, C. phongnhakebangensis, C. calamei by dorsal colour pattern and the number of cloacal and femoral pores in males. However, the new species can be distinguished from C. darevskii by having fewer cloacal and femoral pores in females (maximum 0–28 versus 24–34), four to six blackish brown transverse body bands, as wide as light bands (versus four to five dark irregular transverse breaking body bands, 0.5 times narrower than light band), first body band wide, butterfly-shaped (versus thin, U-shaped in C. darevskii), the presence of tubercles on fore limbs (versus absent), and tail consisting of light rings (versus banded); Cyrtodactylus hinnamnoensis sp. nov. differs from C. phongnhakebangensis by its slightly larger size (SVL reaching 100.6 mm versus 96.3 mm), having fewer cloacal and femoral pores in females (0–28 versus 0–41), having more scale rows from mental to the front of cloacal slit (179–201 versus 161–177), the presence of tubercles on fore limbs (versus absent), a narrower nuchal loop, not enlarged posteriorly (wide, enlarged posteriorly), four to six blackish brown transverse body bands as wide as light bands (versus three to five dark transverse body bands as wide as double light bands, light transverse bands with small spots (versus with big black blotches), and tail pattern consisting of light rings (versus banded); Cyrtodactylus hinnamnoensis sp. nov. differs from C. calamei by its larger size (SVL reaching 100.6 mm versus 89.3 mm), fewer cloacal and femoral pores in females (0–28 versus 38), more postcloacal tubercles (4–6 versus 4), dorsal head marking with distinctly dark spots and blotches (versus indistinct dots), the absence of heart-shaped marking on postocciput (versus present), four to six blackish brown body = standard deviatiοn, fοr οther abbreviatiοns see material and methοds). ……continued on the next page transverse bands, as wide as light bands (versus four greyish brown transverse bands, narrower than light bands) (for more details see Table 7, Fig. 9). The results of the correspondence analysis comparing all adult male morphological maesurements of Cyrtodactylus hinnamnoensis sp. nov. and the latter species indicated four distinct groups between these species (see Fig. 4). Distribution. Cyrtodactylus hinnamnoensis sp. nov. is currently known only from the type locality in the karst forest of Hin Nam No NPA, Khammouane Province, central Laos (Fig. 6). Etymology. We name this species after its type locality, Hin Nam No NPA where the new Cyrtodactylus species was discovered and propose the following common names: Hinnamno Bent-toed Gecko (English), Ki Chiem Hin Nam No (Laotian). Natural history. Specimens were found at night between 19: 41 and 22:03h on karst walls, ca. 0.3–5 m above the ground, near cave entrances in the limestone forest, at elevations between 175 and 580 m a.s.l. Only one male specimen VNUF R. 2014.99 was collected on a tree trunk, about 1 m from the forest floor. The surrounding habitat was karst forest, dominated by species of Ebenaceae, Dracaenaceae, Arecaeae, Poaceae, Meliaceae, and Moraceae. The relative humidity ranged from 78 % to 90 %, and temperatures were from 24.9 to 30.7 o C (see Table 8). When capturing individuals of the species, we observed an increased rate of tail autotomy and many individuals had regenerated tail, for example, seven of 11 specimens of Cyrtodactylus hinnamnoensis sp. nov. had dropped or/and regenerated tails. This suggests that these populations might be under the stress of predators (see also Grismer et al. 2016).

Cyrtodactylus phongnhakebangensis group Species. C. bansocensis, C. calamei, C. darevskii, C. hinnamnoensis, C. jaegeri, C. jarujini, C. khammouanensis, C. lomyenensis, C. multiporus, C. pageli, C. rufford, C. sommerladi, C. soudthichaki, C. teyniei. Characters. Adult SVL 73���100.6 mm, supranasals 0���1; dorsal tubercles 10���24 (except for C. sommerladi, which only has 0���5); webbing between fingers and toes absent; tubercles on fore limbs lacking (except for C. calamei, C. hinnamnoensis, C. jaegeri, C. jarujini, C. multiporus, and C. soudthichaki); tubercles on hind limbs present (except for C. pageli); precloacal and femoral pores in males 20���60 (except for C. pageli only four and for C. teyniei such data is lacking); postcloacal tubercles 3���8 (rarely two); subcaudals enlarged; body bearing welldefined bands 3���8 (except for C. jarujini and C. teyniei, which are blotched). Distribution. The group is mainly distributed in the karst forest mountains of Khammouane Province in central Laos, C. calamei, C. hinnamnoensis, and C. sommerladi are restricted to Hin Nam No NPA in the southeastern Khammouane Province. C. jarujini and C. teyniei are recorded from Bolikhamxay Province, while C. pageli is only known to occur in Vientiane Province. Remarks. The Cyrtodactylus phongnhakebangensis group is very complex. Different morphological traits of the species of this group (e.g., body size, shape and number of dorsal bands, and number of precloacal and femoral pores, see Tables 7 & 13) were supported by other morphological and phylogenetic analyses (see Figs. 1���5). All species of the group are adapted to karst forested formations., Published as part of Luu, Vinh Quang, Bonkowski, Michael, Nguyen, Truong Quang, Le, Minh Duc, Schneider, Nicole, Ngo, Hanh Thi & Ziegler, Thomas, 2016, Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos, pp. 101-140 in Zootaxa 4107 (2) on page 132, DOI: 10.11646/zootaxa.4107.2.1, http://zenodo.org/record/266526

Cyrtodactylus species groups in Laos As currently recognized, 21 Cyrtodactylus species are known from Laos. Based on our molecular phylogeny, the molecular data presented in Nazarov et al. (2014), and a combination of characters including size, dorsal color pattern, scalation, and geographic distribution, we herein propose four species groups within the genus Cyrtodactylus occurring in Laos (see Table 13), which are defined as follows

Cyrtodactylus interdigitalis group Species. C. interdigitalis. Characters. Adult SVL 80.0 mm, supranasal single; dorsal tubercle rows 18; webbing between fingers and toes present; tubercles on limbs present; number of precoacal and femoral pores in males 16���18; postcloacal tubercles 1���2; tail dorsally with tubercles; without enlarged subcaudals; dorsal body blotched or ambiguous. Distribution. The species is known to occur in central Laos including Vientiane and Khammouane provinces. It is adapted as a tree-dweller, but in contrast to the C. irregularis species group that lives near the base of trees, C. interdigitalis is found higher on tree trunks and in tree crowns., Published as part of Luu, Vinh Quang, Bonkowski, Michael, Nguyen, Truong Quang, Le, Minh Duc, Schneider, Nicole, Ngo, Hanh Thi & Ziegler, Thomas, 2016, Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos, pp. 101-140 in Zootaxa 4107 (2) on page 135, DOI: 10.11646/zootaxa.4107.2.1, http://zenodo.org/record/266526