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A new species of the genus Tylototriton is described from Ngoc Linh Mountain, Kon Tum Province, in the Central Highlands of Vietnam based on integrative taxonomy, namely by combining molecular and morphological evidence. Tylototriton ngoclinhensis sp. nov. differs from all other congeners based on morphological data, allopatric distribution, and molecular divergence. In terms of genetic divergence, Tylototriton ngoclinhensis sp. nov. distinctly differs from the sister species T. panhai (6.77%) and from T. ngarsuensis (12.36%) based on the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene. Tylototriton ngoclinhensis sp. nov. is a moderate sized and robust salamander species with large cephalic edges, parotoids, and vertebral ridge orange in coloration. The new taxon differs from its congeners by a combination of the following morphological characteristics: size medium (SVL 60.8–66.5 mm, TL 57.6–61.8 mm in males, and SVL 72.5–75.6 mm, TL 62.9–67.9 mm in females); head longer than wide; parotoids very prominent and enlarged, projecting backwards; tail length shorter than snout-vent length; vertebral ridge large, high and glandular in appearance; 14 large and distinct dorsolateral glandular warts; gular fold present; tips of fore and hind limbs overlapping when adpressed along the body; tips of fingers reaching between eye and nostril when foreleg is laid forward; dorsal surface and lateral sides of the head, upper and lower lips, dorsolateral glandular warts, vertebral ridge, the peripheral area of the cloaca and the ventral edge of the tail orange in coloration; the presence of a distinct black line extending from the posterior end of the eye towards the shoulder. Tylototriton ngoclinhensis sp. nov. is restricted to evergreen montane forests near water bodies on Ngoc Linh Mountain. We suggest that the new species should be classified as Endangered (EN) in the IUCN Red List. This new important discovery represents the eighth Tylototriton taxon described from Vietnam, and at the same time constitutes the southernmost distributional record for the whole genus in Asia.

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

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.

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

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.

Microhyla xodangorum sp. nov. urn:lsid:zoobank.org:act: 2C2DD014-C1A0-439B-8B15-CFB30D715149 Figs 1–3, 5–6, 7B, Tables 1–3, 5 Diagnosis Microhyla xodangorum sp. nov. is distinguished from its congeners by a combination of the following morphological characteristics: 1) size large (SVL 25.6 in a single ♀); 2) snout profile round; 3) dorsal skin almost smooth with flat tubercles evenly scattered in dorsum; dorsolateral edges sharp; 4) chest and belly yellowish with indistinct greyish mottling laterally; chin grey with thickly greyish mottling and a thin, light-coloured medial stripe not reaching the chest; 5) first finger longer than one half of second finger; 6) disks on distal end of fingers II–IV weak; 7) tibiotarsal articulation at straightened limb well beyond snout. Etymology The specific name is a patronym for the Xo Dang people, an ethnic minority people in Kon Tum Province, Vietnam. Their assistance made it possible for us to collect the holotype of the new species in the montane forest of the Kon Tum Plateau, Central Highlands of Vietnam. We recommend ‘Xodang Narrow-mouth Frog’ as the common English name and ‘Nhái bầu xƠ đăng’ as the Vietnamese name. Material examined Holotype VIETNAM • adult ♀; Central Vietnam, Kon Tum Province, Kon Plong District, Mang Canh Commune (Fig. 6); 14°38'49.2″ N, 108°15′39.9″ E; ca 1253m a.s.l.; 15 Sep. 2018; C.V. Hoang leg.; IEBR A.4913 (KP-MĐ-2018-21). Description of holotype (Fig. 6) Preserved specimens were in good condition. HABITUS. Stocky, size medium (SVL 25.6 mm); head wider than long (HL/ HW 0.83); snout long, abruptly round in dorsal view, projecting beyond margin of lower jaw, longer than diameter of eye (SL / EL 1.29); eyes small, slightly protuberant, pupil round (Fig. 6A); nostril oval, lateral, nostril-eyelid length (N-EL 1.4 mm) shorter than eye length (EL 2.2 mm); dorsal surface of head flat, loreal region acute, canthus rostralis round; interorbital distance greater than internarial distance (IOD / IND 1.31) and upper eyelid width (IND / UEW 1.68); tympanum hidden, supratympanic fold weak, from posterior corner of eye to arm insertion; vomerine teeth absent, tongue without papillae, oval and free at rear half of its length. FORELIMBS. Short, about one-half of snout-vent length (FLL / SVL 0.52); hand two times shorter than forelimb length (HAL / FLL 0.45); fingers slender, free of webbing, round in cross-section, dermal fringes weak; first finger longer than one-half length of second finger (1 FLO / 2 FLO 0.53); second finger slightly longer than fourth (2 FLI / 4 FLI 1.35) and shorter than third (2 FLI / 3 FLI 0.67); relative finger lengths: IColoration of holotype in life (Fig. 6A–B) Dorsal surface of head and trunk beige with dark brown dust, forming the markings; a dark brown marking triangle shape between eyelids, continuing and then extending to the groin as a faint dark brown hourglass shape; the biggest black circle ‘o’ shape between middle of hourglass shape; few tiny black scapular spots and black dots scattered around the hourglass shape. Flanks and lateral head dark brown, supratympanic fold and armpit region beige; upper jaw dark brown with few tiny beige spots below the eye level; dorsal surfaces of limbs beige with dark brown crossbars and few small black spots (Fig. 6A). Chest and belly yellowish with indistinct greyish mottling laterally. Chin grey with thickly greyish mottling and a thin, light-coloured medial stripe not reaching the chest (Fig. 6B). Iris golden in upper one-third with black reticulation, dark copper in its lower two-third, darkly pigmented at anterior and posterior corners; pupil round, black (Fig. 6A). Coloration of holotype in preservative After preservation in ethanol, dorsal coloration changed from beige to whitish grey; ventral surface of chest, belly, and limbs changed from dark-beige to dark grey; dorsal pattern, dark spots on dorsum and stripes on dorsal surfaces of limbs unchanged, dark brown pattern changed to dark grey; iris completely black, pupil round, white. Comparisons Microhyla xodangorum sp. nov. differs from other species of the M. heymonsi group by the following characteristics: 1) size large (SVL 25.6 in a ♀) versus Microhyla hmongorum sp. nov. (SVL 13.8–17.4 in 5 ♂♂; 19.2–20.3 in 3 ♀♀), M. ninhthuanensis (SVL 17.3–18.8 in ♂♂; 21.6–23.6 in ♀♀), versus M. pineticola (SVL 17.2–19.5 in ♂♂; 18.0–23.0 in ♀♀); 2) snout profile round versus Microhyla hmongorum, M. pineticola, M. neglecta, (snout profile acuminate) and M. heymonsi s. str. (snout profile obtusely-pointed); 3) dorsum skin almost smooth with flat tubercles evenly scattered in dorsum; dorsolateral edges sharp versus Microhyla hmongorum (smooth with tiny and flat tubercles unevenly scattered, dorsolateral edges not sharp), M. ninhthuanensis (smooth, flanks shagreened; dorsolateral edges not sharp), M. daklakensis (smooth, flanks smoothly shagreened; dorsolateral edges not sharp), M. heymonsi s. str. (smooth, dorsolateral edges not sharp), M. pineticola (almost smooth above with few tiny tubercles scattered in posterior part of dorsum and along the dorsolateral edges), M. neglecta (smooth with evenly scattered small flat tubercles); 4) chest and belly yellowish with indistinct greyish mottling laterally; chin grey with thickly greyish mottling and a thin, light-coloured medial stripe not reaching the chest versus Microhyla hmongorum (chest and belly creamy white fading towards the groin and thighs with indistinct grayish mottling along the thighs and belly edges, chin, throat pinkish white with scattered indistinct grayish mottling), M. ninhthuanensis, M. daklakensis, M. heymonsi s. str. (chest and belly creamy white; chin dark grey; throat white with scattered dark grey dusting), M. pineticola (belly purplish-grey with indistinct whitish mottling; chin dark greyish with orange speckles and a thin light-coloured medial stripe continuing to chest and belly), M. neglecta (chest and belly yellowish with indistinct greyish marbling laterally; centre of chin grey, sides dark brown to black with a thin, lightcoloured medial stripe not reaching the chest); 5) first finger longer than one half of second finger versus M. ninhthuanensis, M. pineticola, M. heymonsi s. str. group (first finger shorter than one half of second finger); 6) disks on distal end of fingers II– IV weak versus Microhyla hmongorum, M. daklakensis, M. ninhthuanensis, M. heymonsi s. str. (disks on distal end of all fingers); 7) tibiotarsal articulation at straightened limb well beyond snout versus Microhyla hmongorum, M. daklakensis, M. ninhthuanensis, M. heymonsi s. str. (shorter than snout). Detailed comparisons between Microhyla xodangorum and other species of the M. heymonsi group are shown in Table 5 and of the genus Microhyla in total are shown in Appendix 1. Natural history All specimens were collected at night from 19:00 to 23:00 on the ground near the banks of a small stream in dipterocarp forest (Fig. 7B). Air temperatue was 21°C and relative humidity was 97%. Larval stages, eggs and the advertisement call of the new species are unknown. Distribution Microhyla xodangorum sp. nov. is currently only known from the type locality in Kon Tum Plateau, Kon Tum Province, Vietnam (Fig. 7 B). The species was found at an elevation of ca 1250 m a.s.l. Based on its habitat and altitudinal range, the new species is likely endemic to the Central Highlands of Vietnam. However, the extent of its actual distribution range requires further studies., Published as part of Van Hoang, Chung, Nguyen, Tao Thien, Phan, Tien Quang, Pham, Cuong The, Ninh, Hoa Thi, Wang, Bin, Jiang, Jianping, Ziegler, Thomas & Nguyen, Truong Quang, 2022, Distribution pattern of the Microhyla heymonsi group (Anura, Microhylidae) with descriptions of two new species from Vietnam, pp. 1-41 in European Journal of Taxonomy 846 on pages 24-27, DOI: 10.5852/ejt.2022.846.1961, http://zenodo.org/record/7268704

Microhyla hmongorum sp. nov. urn:lsid:zoobank.org:act: 3158A3D2-A0D7-43C5-B7E3-C7247567E1AA Figs 2–3, Tables 1–3 Diagnosis Microhyla hmongorum sp. nov. is distinguished from its congeners by a combination of the following morphological characters: 1) size small (SVL 13.8–17.4 in 5 ♂♂; 19.2–20.3 in 3 ♀♀); 2) snout profile acuminate; 3) dorsal skin smooth with tiny and flat tubercles unevenly scattered, dorsolateral edges not sharp; 4) chest and belly creamy-white fades towards the groin and thighs with indistinct grayish mottling along the thighs and belly edges, chin, throat pinkish white with scattered indistinct grayish mottling; 5) first finger longer than one half of second finger. Etymology The specific name is a patronym for the H’Mong people, an ethnic minority people in the northwest montane regions of Vietnam. Their assistance made it possible for us to collect the type specimens of the new species in the montane forest of Lai Chau Province, northwestern Vietnam. We recommend ‘Hmong Narrow-Mouth Frog’ as the common English name and ‘Nhái bầu hmông’ as the Vietnamese name. Material examined Holotype VIETNAM • adult ♂; northern Vietnam, Lai Chau Province, Tam Duong District (Fig. 4); 22°21′15.7″ N, 103°36′36.4″ E; 1362 m a.s.l.; 22 May 2020; C.V. Hoang et al. leg.; IEBR A.4905 (TD-LC2020.121). Paratypes VIETNAM • 4 adult ♂♂; same collection data as for holotype; IEBR A.4906–4907 (TD-LC2020.122, TD-LC2020.123), IEBR A.4908–4909 (TD-LC2020.129, TD-LC2020.130) • 3 adult ♀♀; same collection data as for holotype; IEBR A.4910 (TD-LC2020.124), IEBR A.4911–4912 (TD-LC2020.131, TD-LC2020.132). Description of holotype (Fig. 4, Table 3) Preserved specimens were in good condition. HABITUS. Stocky, SVL 17.4 mm; body triangular shaped; head as long as wide (HL/ HW 1.00); snout long, abruptly round in dorsal view, projecting beyond margin of lower jaw, equal to diameter of eye (SL/ EL 1.02); eyes small, slightly protuberant, pupil round (Fig. 4F); dorsal surface of head flat, canthus rostralis round; loreal region steep, weakly concave; nostril round, lateral, below canthus rostralis, nostril-eyelid length (N-EL 1.3 mm) greater than one half of eye length (N-EL / EL 0.61); interorbital distance (IOD 1.8 mm) greater than internarial distance (IND 1.5 mm) and upper eyelid width (UEW 1.4 mm); pineal spot absent, tympanum hidden, supratympanic fold weak, extending from posterior corner of eye to arm insertion; vomerine teeth absent; tongue without papillae, roundly spatulate and free at the rear half of its length; slit-like openings to a median vocal sac. FORELIMBS. Short, about two times as long as snout–vent length (FLL / SVL 0.57); hand length two times shorter than forelimb length (HAL / FLL 0.42); fingers slender, free of webbing, a little flat in cross-section, with skin fringes on fingers present, dorsoventrally flattened; first finger well developed, longer than one-half length of second finger (1 FLO / 2 FLO 0.69), second finger slightly longer than fourth (2 FLI / 4 FLI 1.32) and longer than one-half length of third finger (2 FLI / 3 FLI 0.68); relative finger lengths: IMicrohyla hmongorum sp. nov. is distinguished from other members of the Microhyla heymonsi group by the following morphological characteristics: 1) body size small (SVL 13.8–17.4 in 5 ♂♂; 19.2–20.3 in 3 ♀♀) versus M. ninhthuanensis (SVL 17.3–18.8 in ♂♂; 21.6–23.6 in ♀♀), M. daklakensis (SVL 17.7– 20.1 in ♂♂; 22.9–26.8 in ♀♀), M. neglecta (SVL 18.7–20.2 in ♂♂; 23.4–26.2 in ♀♀); 2) snout profile acuminate versus M. ninhthuanensis, M. daklakensis (snout profile rounded) and M. heymonsi s. str. (snout profile obtusely pointed); 3) dorsal skin smooth with tiny and flat tubercles unevenly scattered, dorsolateral edges not sharp versus M. ninhthuanensis (smooth, flanks shagreened; dorsolateral edges not sharp), M. daklakensis (smooth, flanks smoothly shagreened; dorsolateral edges not sharp), M. heymonsi s. str. (smooth, dorsolateral edges not sharp), M. pineticola (almost smooth above, with few tiny tubercles scattered in posterior part of dorsum and along the dorsolateral edges; dorsolateral edges sharp), M. neglecta (smooth with evenly scattered small flat tubercles; dorsolateral edges sharp); 4) chest and belly creamy white fades towards the groin and thighs with indistinct grayish mottling along the thighs and belly edges, chin, throat pinkish white with scattered indistinct grayish mottling versus M. ninhthuanensis, M. daklakensis, M. heymonsi s. str. (chest and belly creamy white; chin dark grey; throat white with scattered dark grey dusting), M. pineticola (belly purplish-grey with indistinct whitish mottling; chin dark greyish with orange speckles and a thin light coloured medial stripe continuing to chest and belly), M. neglecta (chest and belly yellowish with indistinct greyish marbling laterally; centre of chin grey, sides dark brown to black with a thin, light-coloured medial stripe not reaching the chest); 5) first finger longer than one half of second finger versus M. ninhthuanensis, M. pineticola, M. heymonsi s. str. (first finger shorter than one half of second finger). Detailed comparisons between Microhyla hmongorum and other species of the M. heymonsi group are shown in Table 5 and of the genus Microhyla in total are shown in Appendix 1. Natural history All specimens were collected at night from 19:00 to 23:00 on the forest trails near streams in the evergreen forest at an elevation of 1300 m a.s.l. (Fig. 7A). Air temperature was 23°C and relative humidity was 89%. Other anuran species found at the type locality were Leptobrachella ventripunctata (Fei, Ye & Li, 1990) and Limnonectes bannaensis Ye, Fei, Xie & Jiang, 2007. Larval stages, eggs and advertisement call of the new species are unknown. Distribution Microhyla hmongorum sp. nov. is known from Hoang Lien Range, NW Vietnam. Garg et al. (2019) reported this species from Phongsali and Luang Prabang provinces of Laos and Poyarkov et al. (2019) recorded the species from Kachin of Myanmar. It is expected to be found in southwestern Yunnan Province, China as well (Fig. 5)., Published as part of Van Hoang, Chung, Nguyen, Tao Thien, Phan, Tien Quang, Pham, Cuong The, Ninh, Hoa Thi, Wang, Bin, Jiang, Jianping, Ziegler, Thomas & Nguyen, Truong Quang, 2022, Distribution pattern of the Microhyla heymonsi group (Anura, Microhylidae) with descriptions of two new species from Vietnam, pp. 1-41 in European Journal of Taxonomy 846 on pages 18-24, DOI: 10.5852/ejt.2022.846.1961, http://zenodo.org/record/7268704, {"references":["Hoang V. C., Nguyen T. T., Ninh T. H., Luong M. A., Pham T. C., Nguyen Q. T., Orlov N., Chen Y. H., Wang B., Ziegler T. & Jiang J. P. 2021. Two new cryptic species of Microhyla Tschudi, 1838 (Amphibia, Anura, Microhylidae) related to the M. heymonsi group from central Vietnam. ZooKeys 1036: 47 - 74. https: // doi. org / 10.3897 / zookeys. 1036.56919","Garg S., Suyesh R., Das A., Jiang J. P., Wijayathilaka N., Amarasinghe A. A. T., Alhadi F., Vineeth K. K., Aravind N. A., Senevirathne G., Meegaskumbura M. & Biju S. D. 2019. Systematic revision of Microhyla","Poyarkov N. A., Gorin V. A., Zaw T., Kretova V. D., Gogoleva S. I., Pawangkhanant P. & Che J. 2019. On the road to Mandalay: contribution to the Microhyla Tschudi, 1838 (Amphibia: Anura: Microhylidae) fauna of Myanmar with description of two new species. Zoological Research 40 (4): 244 - 276. https: // doi. org / 10.24272 / j. issn. 2095 - 8137.2019.044"]}

Microhyla xodangorum sp. nov. urn:lsid:zoobank.org:act: 2C2DD014-C1A0-439B-8B15-CFB30D715149 Figs 1–3, 5–6, 7B, Tables 1–3, 5 Diagnosis Microhyla xodangorum sp. nov. is distinguished from its congeners by a combination of the following morphological characteristics: 1) size large (SVL 25.6 in a single ♀); 2) snout profile round; 3) dorsal skin almost smooth with flat tubercles evenly scattered in dorsum; dorsolateral edges sharp; 4) chest and belly yellowish with indistinct greyish mottling laterally; chin grey with thickly greyish mottling and a thin, light-coloured medial stripe not reaching the chest; 5) first finger longer than one half of second finger; 6) disks on distal end of fingers II–IV weak; 7) tibiotarsal articulation at straightened limb well beyond snout. Etymology The specific name is a patronym for the Xo Dang people, an ethnic minority people in Kon Tum Province, Vietnam. Their assistance made it possible for us to collect the holotype of the new species in the montane forest of the Kon Tum Plateau, Central Highlands of Vietnam. We recommend ‘Xodang Narrow-mouth Frog’ as the common English name and ‘Nhái bầu xƠ đăng’ as the Vietnamese name. Material examined Holotype VIETNAM • adult ♀; Central Vietnam, Kon Tum Province, Kon Plong District, Mang Canh Commune (Fig. 6); 14°38'49.2″ N, 108°15′39.9″ E; ca 1253m a.s.l.; 15 Sep. 2018; C.V. Hoang leg.; IEBR A.4913 (KP-MĐ-2018-21). Description of holotype (Fig. 6) Preserved specimens were in good condition. HABITUS. Stocky, size medium (SVL 25.6 mm); head wider than long (HL/ HW 0.83); snout long, abruptly round in dorsal view, projecting beyond margin of lower jaw, longer than diameter of eye (SL / EL 1.29); eyes small, slightly protuberant, pupil round (Fig. 6A); nostril oval, lateral, nostril-eyelid length (N-EL 1.4 mm) shorter than eye length (EL 2.2 mm); dorsal surface of head flat, loreal region acute, canthus rostralis round; interorbital distance greater than internarial distance (IOD / IND 1.31) and upper eyelid width (IND / UEW 1.68); tympanum hidden, supratympanic fold weak, from posterior corner of eye to arm insertion; vomerine teeth absent, tongue without papillae, oval and free at rear half of its length. FORELIMBS. Short, about one-half of snout-vent length (FLL / SVL 0.52); hand two times shorter than forelimb length (HAL / FLL 0.45); fingers slender, free of webbing, round in cross-section, dermal fringes weak; first finger longer than one-half length of second finger (1 FLO / 2 FLO 0.53); second finger slightly longer than fourth (2 FLI / 4 FLI 1.35) and shorter than third (2 FLI / 3 FLI 0.67); relative finger lengths: I HINDLIMBS. Slender and long (HLL 48.3 mm), tibia length longer than half of snout–vent length (TL/ SVL 0.60); tibiotarsal articulation at straightened limb well beyond snout; foot longer than tibia (FL /TL 1.38); relative toe lengths: I SKIN. Dorsal surface of head and body almost smooth with flat tubercles evenly scattered on dorsum; dorsolateral edges sharp; dorsal surface of hind limbs and cloacal region with few low tubercles scattered; ventral surfaces smooth, present few tubercles in belly face of thighs near cloacal region. Coloration of holotype in life (Fig. 6A–B) Dorsal surface of head and trunk beige with dark brown dust, forming the markings; a dark brown marking triangle shape between eyelids, continuing and then extending to the groin as a faint dark brown hourglass shape; the biggest black circle ‘o’ shape between middle of hourglass shape; few tiny black scapular spots and black dots scattered around the hourglass shape. Flanks and lateral head dark brown, supratympanic fold and armpit region beige; upper jaw dark brown with few tiny beige spots below the eye level; dorsal surfaces of limbs beige with dark brown crossbars and few small black spots (Fig. 6A). Chest and belly yellowish with indistinct greyish mottling laterally. Chin grey with thickly greyish mottling and a thin, light-coloured medial stripe not reaching the chest (Fig. 6B). Iris golden in upper one-third with black reticulation, dark copper in its lower two-third, darkly pigmented at anterior and posterior corners; pupil round, black (Fig. 6A). Coloration of holotype in preservative After preservation in ethanol, dorsal coloration changed from beige to whitish grey; ventral surface of chest, belly, and limbs changed from dark-beige to dark grey; dorsal pattern, dark spots on dorsum and stripes on dorsal surfaces of limbs unchanged, dark brown pattern changed to dark grey; iris completely black, pupil round, white. Comparisons Microhyla xodangorum sp. nov. differs from other species of the M. heymonsi group by the following characteristics: 1) size large (SVL 25.6 in a ♀) versus Microhyla hmongorum sp. nov. (SVL 13.8–17.4 in 5 ♂♂; 19.2–20.3 in 3 ♀♀), M. ninhthuanensis (SVL 17.3–18.8 in ♂♂; 21.6–23.6 in ♀♀), versus M. pineticola (SVL 17.2–19.5 in ♂♂; 18.0–23.0 in ♀♀); 2) snout profile round versus Microhyla hmongorum, M. pineticola, M. neglecta, (snout profile acuminate) and M. heymonsi s. str. (snout profile obtusely-pointed); 3) dorsum skin almost smooth with flat tubercles evenly scattered in dorsum; dorsolateral edges sharp versus Microhyla hmongorum (smooth with tiny and flat tubercles unevenly scattered, dorsolateral edges not sharp), M. ninhthuanensis (smooth, flanks shagreened; dorsolateral edges not sharp), M. daklakensis (smooth, flanks smoothly shagreened; dorsolateral edges not sharp), M. heymonsi s. str. (smooth, dorsolateral edges not sharp), M. pineticola (almost smooth above with few tiny tubercles scattered in posterior part of dorsum and along the dorsolateral edges), M. neglecta (smooth with evenly scattered small flat tubercles); 4) chest and belly yellowish with indistinct greyish mottling laterally; chin grey with thickly greyish mottling and a thin, light-coloured medial stripe not reaching the chest versus Microhyla hmongorum (chest and belly creamy white fading towards the groin and thighs with indistinct grayish mottling along the thighs and belly edges, chin, throat pinkish white with scattered indistinct grayish mottling), M. ninhthuanensis, M. daklakensis, M. heymonsi s. str. (chest and belly creamy white; chin dark grey; throat white with scattered dark grey dusting), M. pineticola (belly purplish-grey with indistinct whitish mottling; chin dark greyish with orange speckles and a thin light-coloured medial stripe continuing to chest and belly), M. neglecta (chest and belly yellowish with indistinct greyish marbling laterally; centre of chin grey, sides dark brown to black with a thin, lightcoloured medial stripe not reaching the chest); 5) first finger longer than one half of second finger versus M. ninhthuanensis, M. pineticola, M. heymonsi s. str. group (first finger shorter than one half of second finger); 6) disks on distal end of fingers II– IV weak versus Microhyla hmongorum, M. daklakensis, M. ninhthuanensis, M. heymonsi s. str. (disks on distal end of all fingers); 7) tibiotarsal articulation at straightened limb well beyond snout versus Microhyla hmongorum, M. daklakensis, M. ninhthuanensis, M. heymonsi s. str. (shorter than snout). Detailed comparisons between Microhyla xodangorum and other species of the M. heymonsi group are shown in Table 5 and of the genus Microhyla in total are shown in Appendix 1. Natural history All specimens were collected at night from 19:00 to 23:00 on the ground near the banks of a small stream in dipterocarp forest (Fig. 7B). Air temperatue was 21°C and relative humidity was 97%. Larval stages, eggs and the advertisement call of the new species are unknown. Distribution Microhyla xodangorum sp. nov. is currently only known from the type locality in Kon Tum Plateau, Kon Tum Province, Vietnam (Fig. 7 B). The species was found at an elevation of ca 1250 m a.s.l. Based on its habitat and altitudinal range, the new species is likely endemic to the Central Highlands of Vietnam. However, the extent of its actual distribution range requires further studies.

Microhyla hmongorum sp. nov. urn:lsid:zoobank.org:act: 3158A3D2-A0D7-43C5-B7E3-C7247567E1AA Figs 2–3, Tables 1–3 Diagnosis Microhyla hmongorum sp. nov. is distinguished from its congeners by a combination of the following morphological characters: 1) size small (SVL 13.8–17.4 in 5 ♂♂; 19.2–20.3 in 3 ♀♀); 2) snout profile acuminate; 3) dorsal skin smooth with tiny and flat tubercles unevenly scattered, dorsolateral edges not sharp; 4) chest and belly creamy-white fades towards the groin and thighs with indistinct grayish mottling along the thighs and belly edges, chin, throat pinkish white with scattered indistinct grayish mottling; 5) first finger longer than one half of second finger. Etymology The specific name is a patronym for the H’Mong people, an ethnic minority people in the northwest montane regions of Vietnam. Their assistance made it possible for us to collect the type specimens of the new species in the montane forest of Lai Chau Province, northwestern Vietnam. We recommend ‘Hmong Narrow-Mouth Frog’ as the common English name and ‘Nhái bầu hmông’ as the Vietnamese name. Material examined Holotype VIETNAM • adult ♂; northern Vietnam, Lai Chau Province, Tam Duong District (Fig. 4); 22°21′15.7″ N, 103°36′36.4″ E; 1362 m a.s.l.; 22 May 2020; C.V. Hoang et al. leg.; IEBR A.4905 (TD-LC2020.121). Paratypes VIETNAM • 4 adult ♂♂; same collection data as for holotype; IEBR A.4906–4907 (TD-LC2020.122, TD-LC2020.123), IEBR A.4908–4909 (TD-LC2020.129, TD-LC2020.130) • 3 adult ♀♀; same collection data as for holotype; IEBR A.4910 (TD-LC2020.124), IEBR A.4911–4912 (TD-LC2020.131, TD-LC2020.132). Description of holotype (Fig. 4, Table 3) Preserved specimens were in good condition. HABITUS. Stocky, SVL 17.4 mm; body triangular shaped; head as long as wide (HL/ HW 1.00); snout long, abruptly round in dorsal view, projecting beyond margin of lower jaw, equal to diameter of eye (SL/ EL 1.02); eyes small, slightly protuberant, pupil round (Fig. 4F); dorsal surface of head flat, canthus rostralis round; loreal region steep, weakly concave; nostril round, lateral, below canthus rostralis, nostril-eyelid length (N-EL 1.3 mm) greater than one half of eye length (N-EL / EL 0.61); interorbital distance (IOD 1.8 mm) greater than internarial distance (IND 1.5 mm) and upper eyelid width (UEW 1.4 mm); pineal spot absent, tympanum hidden, supratympanic fold weak, extending from posterior corner of eye to arm insertion; vomerine teeth absent; tongue without papillae, roundly spatulate and free at the rear half of its length; slit-like openings to a median vocal sac. FORELIMBS. Short, about two times as long as snout–vent length (FLL / SVL 0.57); hand length two times shorter than forelimb length (HAL / FLL 0.42); fingers slender, free of webbing, a little flat in cross-section, with skin fringes on fingers present, dorsoventrally flattened; first finger well developed, longer than one-half length of second finger (1 FLO / 2 FLO 0.69), second finger slightly longer than fourth (2 FLI / 4 FLI 1.32) and longer than one-half length of third finger (2 FLI / 3 FLI 0.68); relative finger lengths: I HINDLIMBS. Slender and slightly short (HLL 30.5 mm), tibia length longer than half of snout–vent length (TL/ SVL 0.53); tibiotarsal articulation at straightened limb not reaching snout; foot longer than tibia (FL /TL 1.54); relative toe lengths: I SKIN. Dorsal surface smooth with tiny and flat tubercles unevenly scattered; dorsolateral edges not sharp; upper eyelid without supraciliary spines; flanks of body and lateral sides of head smooth, hindlimb dorsally scattered with some low pustules; ventral side of body and limbs smooth, vent area smooth with several low tubercles in cloacal region (Fig. 4B). Cloacal opening unmodified, directed posteriorly, at lower level of thighs. Coloration of holotype in life Dorsal surface of head and trunk beige with dark brown dust forming blurred markings; two dark brown tiny spots between eyelids; a small dark brown marking in ‘()’-shape in the center of dorsum; a middorsal line extending from the tip of snout to vent; a dark brown rhombus pattern on dorsum: one side is a small dark marking in ‘()’-shape, one side is a small dark brown blurred marking in ‘()’-shape. Flanks and lateral side of head dark with a dark lateral stripe running from tip of snout to nostril, fading towards upper jaw (Fig. 4A). Chest and belly creamy white fading towards the groin and thighs with indistinct grayish mottling along the thighs and belly edges. Chin, throat pinkish white with scattered indistinct grayish mottling (Fig. 4B). Limbs dorsally beige with narrow blurred dark brown crossbars; fingers and toes dorsally brown with scattered dark brown crossbars; limbs ventrally crystal-clear with scattered dark grey and white dust, getting thicker toward shank and foot. Iris bicolored, golden in upper third, dark copper in its lower two thirds; pupil oval, horizontal, black. Coloration of holotype in preservative After preservation in ethanol, dorsal coloration changed from beige to greyish beige; ventral surface of chest, belly, and limbs changed from crystal clear to white; dorsal pattern, dark spots on dorsum and stripes on dorsal surfaces of limbs unchanged, dark brown pattern changed to dark grey; iris completely black, pupil round, white. Variation The paratypes vary in body size, coloration of dorsal surface, dorsal marking and black scapular spots. Adult males are smaller than adult females. The female’s belly is whiter than that of the male. Comparisons Microhyla hmongorum sp. nov. is distinguished from other members of the Microhyla heymonsi group by the following morphological characteristics: 1) body size small (SVL 13.8–17.4 in 5 ♂♂; 19.2–20.3 in 3 ♀♀) versus M. ninhthuanensis (SVL 17.3–18.8 in ♂♂; 21.6–23.6 in ♀♀), M. daklakensis (SVL 17.7– 20.1 in ♂♂; 22.9–26.8 in ♀♀), M. neglecta (SVL 18.7–20.2 in ♂♂; 23.4–26.2 in ♀♀); 2) snout profile acuminate versus M. ninhthuanensis, M. daklakensis (snout profile rounded) and M. heymonsi s. str. (snout profile obtusely pointed); 3) dorsal skin smooth with tiny and flat tubercles unevenly scattered, dorsolateral edges not sharp versus M. ninhthuanensis (smooth, flanks shagreened; dorsolateral edges not sharp), M. daklakensis (smooth, flanks smoothly shagreened; dorsolateral edges not sharp), M. heymonsi s. str. (smooth, dorsolateral edges not sharp), M. pineticola (almost smooth above, with few tiny tubercles scattered in posterior part of dorsum and along the dorsolateral edges; dorsolateral edges sharp), M. neglecta (smooth with evenly scattered small flat tubercles; dorsolateral edges sharp); 4) chest and belly creamy white fades towards the groin and thighs with indistinct grayish mottling along the thighs and belly edges, chin, throat pinkish white with scattered indistinct grayish mottling versus M. ninhthuanensis, M. daklakensis, M. heymonsi s. str. (chest and belly creamy white; chin dark grey; throat white with scattered dark grey dusting), M. pineticola (belly purplish-grey with indistinct whitish mottling; chin dark greyish with orange speckles and a thin light coloured medial stripe continuing to chest and belly), M. neglecta (chest and belly yellowish with indistinct greyish marbling laterally; centre of chin grey, sides dark brown to black with a thin, light-coloured medial stripe not reaching the chest); 5) first finger longer than one half of second finger versus M. ninhthuanensis, M. pineticola, M. heymonsi s. str. (first finger shorter than one half of second finger). Detailed comparisons between Microhyla hmongorum and other species of the M. heymonsi group are shown in Table 5 and of the genus Microhyla in total are shown in Appendix 1. Natural history All specimens were collected at night from 19:00 to 23:00 on the forest trails near streams in the evergreen forest at an elevation of 1300 m a.s.l. (Fig. 7A). Air temperature was 23°C and relative humidity was 89%. Other anuran species found at the type locality were Leptobrachella ventripunctata (Fei, Ye & Li, 1990) and Limnonectes bannaensis Ye, Fei, Xie & Jiang, 2007. Larval stages, eggs and advertisement call of the new species are unknown. Distribution Microhyla hmongorum sp. nov. is known from Hoang Lien Range, NW Vietnam. Garg et al. (2019) reported this species from Phongsali and Luang Prabang provinces of Laos and Poyarkov et al. (2019) recorded the species from Kachin of Myanmar. It is expected to be found in southwestern Yunnan Province, China as well (Fig. 5).

Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tien, Phan Quang, Tran, Hieu Minh, Nguyen, Truong Quang, van Schingen-Khan, Mona, Ziegler, Thomas (2022): Ecological niche overlap of two allopatric karst-adapted tiger geckos (Goniurosaurus) from northern Vietnam: microhabitat use and implications for conservation. Journal of Natural History 56 (37-40): 1495-1511, DOI: 10.1080/00222933.2022.2120437, URL: http://dx.doi.org/10.1080/00222933.2022.2120437

Xenophrys truongsonensis sp. nov. (Figs. 3, 4) Megophrys sp 12., Chen et al. 2017 Megophrys cf. maosoensis 3, Mahony et al. 2018 Holotype. IEBR A.4940, adult male, collected by A.M. Luong, C. V. Hoang, T. V. Nguyen on 24 May 2018 in the evergreen forest (12 o 23’13.8”N, 108 o 20’33.9”E, at an elevation of 1267 m asl.) within Chu Yang Sin National Park, Krong Bong District, Dak Lak Province, Vietnam. Paratypes (n = 18). Specimens collected from the forest of A Luoi District, Thua Thien Hue Province, Vietnam: two adult males IEBR A.4934–4935 (AL.2017.98, 107) and one adult female IEBR A.4936 (AL.2017.106) collected by T. V. Nguyen, C. T. Pham on 6 April 2017 in the evergreen forest (16 o 08’39.32”N, 107 o 35’47.56”E, at an elevation of 713 m asl.); one adult male IEBR A.4937 (AL.2018.1) collected by T.Q. Phan on 16 March 2018 in the evergreen forest (16 o 10’45.43”N, 107 o 29’53.22”E, at an elevation of 798 m asl.). Specimens collected from the forest of Kon Plong District, Kon Tum Province, Vietnam: two adult males IEBR A.4941–4942 (NL.116, 161) and adult female IEBR A.4943 collected by A.M. Luong, V. H.D. Nguyen, N. Orlov on 9 November 2018 in the evergreen forest (14 o 40’30.7”N, 108 o 15’35.1”E, at an elevation of 1319 m asl.). Specimens collected from the forest of Chu Yang Sin National Park, Krong Bong District, Dak Lak Province, Vietnam: one adult male IEBR A.4944 collected by A.M. Luong, C. V. Hoang, T. V. Nguyen on 24 May 2018 in the evergreen forest (12 o 23’13.8”N, 108 o 20’33.9”E, at an elevation of 1267 m asl.); one adult male IEBR A.4938 (CYS.2019.69), collected by A.M. Luong and Q.H. Do on 11 March 2019 (12 o 23’12.7”N, 108 o 20’33.4”E, at an elevation of 1270 m asl.). Specimens collected from the forest of Nam Ka Nature Reserve, Lak District, Dak Lak Province, Vietnam: one adult male IEBR A.4945 collected by A.M. Luong, C. V. Hoang, T. V. Nguyen on 28 May 2018 in the evergreen forest (12 o 20’11.5”N, 108 o 01’51.4”E, at an elevation of 532 m asl.). Specimens collected from the forest of Nam Nung Nature Reserve, Dak Glong District, Dak Nong Province, Vietnam: two adult males IEBR A.4946–4947 collected by A.M. Luong on 23 October 2017 in the evergreen forest (12 o 12’24.21”N, 107 o 50’41.23”E, at an elevation of 794 m asl.). Specimens collected from the forest of Bi Doup-Nui Ba National Park, Lac Duong District, Lam Dong Province, Vietnam: two adult males IEBR A.4948–4949 and two adult females IEBR A.4950–4951 collected by A.M. Luong, C. V. Hoang, Y. T. Nguyen, L. V. Pham, T. D. Pham on 14 May 2018 in the evergreen forest (12 o 05’50.2”N, 108 o 21’25.3”E, at an elevation of 1605 m asl.). Specimens collected from the forest of Phuoc Binh National Park, Phuoc Binh District, Ninh Thuan Province, Vietnam: one adult male IEBR A.4952 and one adult female IEBR A.4953 collected by A.M. Luong, C. V. Hoang on 26 April 2018 in the evergreen forest (12 o 01’26.0”N, 108 o 44’16.9”E, at an elevation of 823 m asl.). Diagnosis. The new species is distinguishable from its congeners by a combination of the following morphological characters: 1) size medium (SVL 58.8–71.4 mm in males; SVL 65.6–87.3 mm in females); 2) tympanum distinct (TD/ED 0.51–0.67 in males; TD/ED 0.53–0.59 in females); 3) vomerine teeth present; 4) tongue feebly notched posteriorly; 5) external vocal sac absent; 6) toes with rudimentary webbing; 7) subarticular tubercles absent; 8) lateral fringes narrow along toes; 9) dorsal skin of body with weak granules; 10) flank with small tubercles; 11) dorsum with a dorsal ridge, in >-< shape; 12) dorsolateral folds prominent; 13) dorsal surface from reddish brown to grayish brown with a brown triangle between the eyes, and a dark brown marking edged along dorsal ridge on the back; and 14) small white spots present on flank and back of thigh. Description of holotype. Adult male; SVL 67.2 mm; head as long as wide (HL 26.2 mm, HW 26.2 mm); snout round in dorsal view, obtusely protruding in lateral view; nostril lateral, oval, closer to the tip of snout than to the eye (NS 3.9 mm, EN 4.5 mm); canthus rostralis angular; pupil vertically elliptical; loreal region acute, concave; eye diameter shorter than snout length (SL 8.4 mm, ED 8.0 mm); internarial distance narrower than interorbital distance but wider than upper eyelid (IND 7.2 mm, IOD 7.6 mm, UEW 5.3mm); tympanum distinct, oval, 51% of eye diameter (TD 4.1 mm); vomerine teeth present, maxillary teeth present; tongue round, feebly notched posteriorly; external vocal sac absent. Forelimbs: Hand length (HAL) 34.1 mm, forearm length (FLL) 16.0 mm; relative finger lengths I>-< shape; dorsolateral folds prominent; supratympanic fold distinct; flank with small tubercles; dorsal surface of limbs granular with short transverse ridges; throat, chest, belly and ventral surface of thigh smooth. Coloration in life: Dorsum reddish brown with a dark brown triangle between the eyes; a dark brown marking edged along dorsal ridge; flanks yellowish brown with some small black spots; tympanum dark brown; pupil black, outlined in reddish brown, iris copper-brown with tiny dark reticulations spreading from pupil; upper lip with a continuous white stripe, running from the nostril to shoulder; small white spots on flank and back of thigh; dorsal surface of fore and hind limbs reddish brown with dark crossbars; throat and chest dark brown, pectoral glands white; anterior part of belly white with brown pattern, posterior part of belly white; ventral surface of limbs dark brown with white marking, anterior part of thigh, posterior part of thigh dark brown; femoral glands white; toe webbing dark brown. Coloration in preservative: Dorsum greyish with a dark brown triangle marking between the eyes; a dark brown marking edged along dorsal ridge; flanks greyish with some small dark spots; throat and chest brown; belly cream with brown pattern; dorsal surface of fore and hind limbs greyish with dark brown crossbars; posterior part of thigh dark brown; femoral gland white; toe webbing dark brown. Variation. Measurements and morphological characters of the type series are given in Table 3. Adult males are smaller than adult females. Dorsum is reddish brown or grayish brown. Comparisons. We compared the new species with other species of Xenophrys from Vietnam and neighboring countries, including Laos, Cambodia, Thailand, and China. Xenophrys truongsonensis sp. nov. can be distinguished from its congeners by morphological characters as shown in Table 4. Morphologically, Xenophrys truongsonensis sp. nov. is most similar to X. maosonensis Bourret but they are different from each other by having flank with small tubercles (vs. flank with large tubercles in X. maosonensis), and toes with narrow lateral fringes (vs. absent in X. maosonensis), and white spots on flank and back of thigh smaller than those in X. maosonensis. Xenophrys truongsonensis sp. nov. differs from X. aceras Boulenger by having a greater TbL/SVL ratio in males (0.55–0.64 vs. 0.42–0.46 in X. aceras), and a greater ratio HL/HW>1 (vs. M. aceras); from X. ancrae Mahony, Teeling & Biju by having a larger body size (SVL 58.8–71.4 mm in males, 65.6–87.3 mm in females vs. 39.1–45.0 in males, 48.9 mm in female in X. ancrae), a greater ratio of TbL/SVL in males (0.55–0.64 vs. 0.48 in X. ancrae), toes with narrow lateral fringes (vs. absent in X. ancrae), and the absence of nuptial pads on finger I (vs. present in X. ancrae); from X. auralensis Ohler, Swan & Daltry by having a greater ratio of TbL/SVL in males (0.55–0.64 vs. 0.51 in X. auralensis), the presence of vomerine teeth (vs. absent in X. auralensis), the absence of external vocal sac (vs. present in X. auralensis), the first finger being shortest (vs. the second finger or fourth finger being shortest in X. auralensis), and tongue feebly notched (vs. tongue not notched in X. auralensis); from X. damrei Mahony by having a greater ratio of TbL/SVL in males (0.55–0.64 vs. 0.51 in X. damrei), the absence of external vocal sac (vs. present in X. damrei), and the first finger being shortest (vs. the fourth finger being shortest in X. damrei); from X. glandulosa Fei, Ye, & Huang by the absence of external vocal sac (vs. present in X. glandulosa), toes with narrow lateral fringes (vs. well developed fringes in X. glandulosa), and the first finger being the shortest (vs. the second finger or fourth finger being shortest in X. glandulosa); from X. lekaguli Stuart, Chuaynkern, Chanard, & Inger by having a greater ratio of TbL/SVL (0.55–0.64 in males and 0.54–0.58 mm in females vs. 0.47–0.52 in males and 0.44–0.52 in females in X. lekaguli), tongue feebly notched (vs. tongue not notched in X. lekaguli), toes with narrow lateral fringes (vs. absent in X. lekaguli), and the first finger being shortest (vs. the fourth finger being shortest in X. lekaguli); from X. longipes Boulenger by having a larger body size in males (SVL 58.8–71.4 mm vs. 47 mm in the male X. longipes), the absence of external vocal sac (vs. present in X. longipes), and the first finger being shortest (vs. the second finger being shortest in X. longipes); from X. mangshanensis Fei & Ye by the absence of external vocal sac (vs. present in X. mangshanensis), toes with narrow lateral fringes (vs. absent in X. mangshanensis), toes with rudiment of webbing (vs. without webbing in X. mangshanensis), and the fisrt finger being shortest (vs. the second finger being shortest in X. mangshanensis); from X. medogensis Fei, Ye, & Huang by the absence of external vocal sac (vs. present in X. medogensis), and toes with narrow lateral fringes (vs. absent in X. medogensis); from X. pachyproctus Huang by having a larger body size (SVL 58.8–71.4 mm in males and 65.6–87.3 mm in females vs. 35.3–36.2 mm in males and 35.8 mm in the female in X. pachyproctus), a greater ratio of TD/ED (0.51–0.67 in males and 0.53–0.59 in females vs. 0.35 in the male and 0.36 in the female in X. pachyproctus), a greater ratio of TbL/SVL (0.55–0.6 in males and 0.54–0.58 mm in females vs. 0.48–0.49 in males and 0.5 in the female in X. pachyproctus), the presence of vomerine teeth (vs. absent in X. pachyproctus), toes with narrow lateral fringes (vs. absent in X. pachyproctus), the absence of subarticular tubercles on toes (vs. present in X. pachyproctus), and the absence of nuptial pads on finger I (vs. present in X. pachyproctus); from X. takensis Mahony by having a larger body size in males (58.8–71.4 mm vs. 47.3–53.0 mm in X. takensis), a greater ratio of TD/ED in males (0.51– 0.67 vs 0.42–0.48 in X. takensis), a greater ratio of TbL/SVL (0.55–0.64 in males and 0.54–0.58 mm in females vs. 0.41–0.49 in males and 0.45 in female in X. takensis), the absence of external vocal sac (vs. present in X. takensis), tongue feebly notched (vs. tongue not notched in X. takensis), toes with narrow lateral fringes (vs. absent in X. takensis), the first finger being shortest (vs. the fourth finger being shortest in X. takensis), and the absence of nuptial pads on finger I (vs. present in X. takensis); from X. yeae by having a larger body size (58.8–71.4 mm in males and 65.6–87.3 mm in females vs. 23.0– 29.1 mm in males and 27.9–31.1 mm in the females in X. yeae), a greater ratio of TD/ED (0.55–0.64 in males and and 0.53–0.59 in females vs. 0.38–0.47 in males and 0.39–0.50 in females in X. yeae), and the presence of vomerine teeth (vs. absent in X. yeae); from X. zhangi Ye & Fei by having a larger body size in males (58.8–71.4 mm vs. 32.5–37.2 mm in X. zhangi), a greater ratio of TD/ED in males (0.51–0.67 vs 0.5 in X. zhangi), a greater ratio of TbL/SVL in males (0.55–0.64 vs. 0.49 in X. zhangi), the absence of external vocal sac (vs. present in X. zhangi), and the absence of nuptial pads on finger I (vs. present in X. zhangi); and from X. zhoui by having a larger body size (58.8–71.4 mm in males and 65.6–87.3 mm in females vs. 23 mm in male and 23.5–23.9 mm in the females in X. zhoui), a greater ratio of TD/ED in males (0.55–0.64 vs. 0.48 in X. zhoui), the presence of vomerine teeth (vs. absent in X. yeae), and toes with narrow lateral fringes (vs. absent in X. zhoui). ......continued on the next page ......continued on the next page Etymology. Specific epithet is in reference to the type locality of the Truong Son Mountain Range. We recommend “Truongson Horned Toad” as the common English name and “Cóc mắt trường sƠn” as the Vietnamese name. Ecological notes. Specimens were collected at night between 19:00 and 23:00 on tree leaves, ca 0.2–0.6 m above the ground, or on ground in rocky streams. The surrounding habitat was secondary evergreen forest consisting of medium and small hardwoods mixed with shrubs or secondary evergreen forest consisting of small hardwoods mixed with shrubs (Fig. 5). Distribution. Xenophrys truongsonensis sp. nov. is currently known from Thua Thien Hue Province southwards to Lam Dong Province, Central Vietnam (Fig. 6). It is expected that this species can be found in the evergreen forests on western side of Truong Son Range in Laos and Cambodia as well., Published as part of Luong, Anh Mai, Hoang, Chung Van, Pham, Cuong The, Nguyen, Tao Thien, Orlov, Nikolai, Ziegler, Thomas & Nguyen, Truong Quang, 2022, A new species of Xenophrys (Amphibia: Anura: Megophryidae) from Truong Son Range, Vietnam, pp. 333-356 in Zootaxa 5150 (3) on pages 341-352, DOI: 10.11646/zootaxa.5150.3.2, http://zenodo.org/record/6622887, {"references":["Chen, J., Zhou, W., Poyarkov, N. A., Stuart, B. L., Brown, R. M., Lathrop, A., Wang, Y., Yuan, Z., Jiang, K., Hou, M., Chen, H., Suwannapoom, C, Nguyen, S. N., Duong, T. V., Papenfuss, T. J., Murphy, R. W., Zhang, Y. & Che, J. (2017) A novel multilocus phylogenetic estimation reveals unrecognized diversity in Asian horned toads, genus Megophrys sensu lato (Anura: Megophryidae). Molecular Phylogenetics and Evolution, 106, 28 - 43. https: // doi. org / 10.1016 / j. ympev. 2016.09.004","Mahony, S., Kamei, R. G., Teeling, E. C. & Biju, S. D. (2018) Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species. Zootaxa, 4523 (1), 1 - 96. https: // doi. org / 10.11646 / zootaxa. 4523.1.1","Boulenger, G. A. (1908) A revision of the oriental pelobatid batrachians (genus Megalophrys). Proceedings of the Zoological Society of London, 1908, 407 - 430. https: // doi. org / 10.1111 / j. 1096 - 3642.1908. tb 01852. x","Bourret, R. (1937) Notes herpetologiques sur l'Indochine francaise. XIV. Les batraciens de la collection du Laboratoire des Sciences Naturelles de l'Universite. Descriptions de quinze especes ou varietes nouvelles. Annexe au Bulletin General de l'Instruction Publique, 1937, 5 - 56.","Bourret, R. (1942) Les Batraciens de l'Indochine. Institut Oceanographique de l'Indochine, Hanoi, 567 pp.","Taylor, E. H. (1962) The amphibian fauna of Thailand. University of Kansas Science Bulletin 43. University of Kansas, Lawrence, Kansas, pp. 265 - 599. https: // doi. org / 10.5962 / bhl. part. 13347","Fei, L., Ye, C. Y. & Huang, Y. (1992) Description of two new species of the genus Megophrys, Pelobatidae (Amphibia: Anura) from China. Zoological Research, 13, 5 - 12.","Huang, C. M., Lathrop, A. & Murphy, R. W. (1998) Translations of \" Two new species of amphibian in Tibet \" Huang & Fei 1981 and \" Description of two new species of the genus Megophrys \" Fei, Ye, & Huang 1995. Smithsonian Herpetological Information Service, 118, 1 - 20.","Ohler, A., Swan, S. R. & Daltry, J. C. (2002) A recent survey of the amphibian fauna of the Cardomom Mountains, Southwest Cambodia with descriptions of three new species. Raffles Bulletin of Zoology, 50 (2), 465 - 481.","Stuart, B. L., Chuaynkern, Y., Chan-ard, T. & Inger, R. F. (2006) Three new species of frogs and a new tadpole from eastern Thailand. Fieldiana, 111, 1 - 19.","Fei, L., Hu, S., Ye, C. & Huang, Y. (2009) Fauna Sinica. Amphibia. Vol. 2. Anura. Chinese Academy of Science, Science Press, Beijing, 957 pp. [in Chinese]","Fei, L., Ye, C. & Jiang, J. (2010) Colored Atlas of Chinese Amphibians. Sichuan Publishing House of Science & Technology, Sichuan, 519 pp. [in Chinese]","Mahony, S., Sengupta, S., Kamei, R. G. & Biju, S. D. (2011) A new low altitude species of Megophrys Kuhl & van Hasselt (Amphibia: Megophryidae), from Assam, Northeast India. Zootaxa, 3059 (1), 36 - 46. https: // doi. org / 10.11646 / zootaxa. 3059.1.2","Mahony, S., Teeling, E. C. & Biju, S. D. (2013) Three new species of horned frogs, Megophrys (Amphibia: Megophryidae), from northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region. Zootaxa, 3722 (2), 143 - 169. https: // doi. org / 10.11646 / zootaxa. 3931.2.3","Neang, T., Chin, S., Meang, M. & Hun, S. (2013) Confirmation of three species of megophryid frogs (Amphibia: Megophryidae) from the Cardamom Mountains of Southwest Cambodia, with the rediscovery of a long lost species. Cambodian Journal of Natural History, 2013 (2), 66 - 72.","Shi, S., Zhang, M., Xie, F., Jiang, J., Liu, W., Ding, L., Luan, L. & Wang, B. (2020) Multiple data revealed two new species of the Asian horned toad Megophrys Kuhl & Van Hasselt, 1822 (Anura, Megophryidae) from the eastern corner of the Himalayas. ZooKeys, 977, 101 - 161 https: // doi. org / 10.3897 / zookeys. 977.55693"]}

Xenophrys truongsonensis sp. nov. (Figs. 3, 4) Megophrys sp 12., Chen et al. 2017 Megophrys cf. maosoensis 3, Mahony et al. 2018 Holotype. IEBR A.4940, adult male, collected by A.M. Luong, C. V. Hoang, T. V. Nguyen on 24 May 2018 in the evergreen forest (12 o 23’13.8”N, 108 o 20’33.9”E, at an elevation of 1267 m asl.) within Chu Yang Sin National Park, Krong Bong District, Dak Lak Province, Vietnam. Paratypes (n = 18). Specimens collected from the forest of A Luoi District, Thua Thien Hue Province, Vietnam: two adult males IEBR A.4934–4935 (AL.2017.98, 107) and one adult female IEBR A.4936 (AL.2017.106) collected by T. V. Nguyen, C. T. Pham on 6 April 2017 in the evergreen forest (16 o 08’39.32”N, 107 o 35’47.56”E, at an elevation of 713 m asl.); one adult male IEBR A.4937 (AL.2018.1) collected by T.Q. Phan on 16 March 2018 in the evergreen forest (16 o 10’45.43”N, 107 o 29’53.22”E, at an elevation of 798 m asl.). Specimens collected from the forest of Kon Plong District, Kon Tum Province, Vietnam: two adult males IEBR A.4941–4942 (NL.116, 161) and adult female IEBR A.4943 collected by A.M. Luong, V. H.D. Nguyen, N. Orlov on 9 November 2018 in the evergreen forest (14 o 40’30.7”N, 108 o 15’35.1”E, at an elevation of 1319 m asl.). Specimens collected from the forest of Chu Yang Sin National Park, Krong Bong District, Dak Lak Province, Vietnam: one adult male IEBR A.4944 collected by A.M. Luong, C. V. Hoang, T. V. Nguyen on 24 May 2018 in the evergreen forest (12 o 23’13.8”N, 108 o 20’33.9”E, at an elevation of 1267 m asl.); one adult male IEBR A.4938 (CYS.2019.69), collected by A.M. Luong and Q.H. Do on 11 March 2019 (12 o 23’12.7”N, 108 o 20’33.4”E, at an elevation of 1270 m asl.). Specimens collected from the forest of Nam Ka Nature Reserve, Lak District, Dak Lak Province, Vietnam: one adult male IEBR A.4945 collected by A.M. Luong, C. V. Hoang, T. V. Nguyen on 28 May 2018 in the evergreen forest (12 o 20’11.5”N, 108 o 01’51.4”E, at an elevation of 532 m asl.). Specimens collected from the forest of Nam Nung Nature Reserve, Dak Glong District, Dak Nong Province, Vietnam: two adult males IEBR A.4946–4947 collected by A.M. Luong on 23 October 2017 in the evergreen forest (12 o 12’24.21”N, 107 o 50’41.23”E, at an elevation of 794 m asl.). Specimens collected from the forest of Bi Doup-Nui Ba National Park, Lac Duong District, Lam Dong Province, Vietnam: two adult males IEBR A.4948–4949 and two adult females IEBR A.4950–4951 collected by A.M. Luong, C. V. Hoang, Y. T. Nguyen, L. V. Pham, T. D. Pham on 14 May 2018 in the evergreen forest (12 o 05’50.2”N, 108 o 21’25.3”E, at an elevation of 1605 m asl.). Specimens collected from the forest of Phuoc Binh National Park, Phuoc Binh District, Ninh Thuan Province, Vietnam: one adult male IEBR A.4952 and one adult female IEBR A.4953 collected by A.M. Luong, C. V. Hoang on 26 April 2018 in the evergreen forest (12 o 01’26.0”N, 108 o 44’16.9”E, at an elevation of 823 m asl.). Diagnosis. The new species is distinguishable from its congeners by a combination of the following morphological characters: 1) size medium (SVL 58.8–71.4 mm in males; SVL 65.6–87.3 mm in females); 2) tympanum distinct (TD/ED 0.51–0.67 in males; TD/ED 0.53–0.59 in females); 3) vomerine teeth present; 4) tongue feebly notched posteriorly; 5) external vocal sac absent; 6) toes with rudimentary webbing; 7) subarticular tubercles absent; 8) lateral fringes narrow along toes; 9) dorsal skin of body with weak granules; 10) flank with small tubercles; 11) dorsum with a dorsal ridge, in >-< shape; 12) dorsolateral folds prominent; 13) dorsal surface from reddish brown to grayish brown with a brown triangle between the eyes, and a dark brown marking edged along dorsal ridge on the back; and 14) small white spots present on flank and back of thigh. Description of holotype. Adult male; SVL 67.2 mm; head as long as wide (HL 26.2 mm, HW 26.2 mm); snout round in dorsal view, obtusely protruding in lateral view; nostril lateral, oval, closer to the tip of snout than to the eye (NS 3.9 mm, EN 4.5 mm); canthus rostralis angular; pupil vertically elliptical; loreal region acute, concave; eye diameter shorter than snout length (SL 8.4 mm, ED 8.0 mm); internarial distance narrower than interorbital distance but wider than upper eyelid (IND 7.2 mm, IOD 7.6 mm, UEW 5.3mm); tympanum distinct, oval, 51% of eye diameter (TD 4.1 mm); vomerine teeth present, maxillary teeth present; tongue round, feebly notched posteriorly; external vocal sac absent. Forelimbs: Hand length (HAL) 34.1 mm, forearm length (FLL) 16.0 mm; relative finger lengths I Hindlimbs: Tibia longer than thigh (FeL 35.9 mm, TbL 38.7 mm), approximately five times longer than wide (TbW 7.7 mm); tips of toes round, relative length of toes I Skin texture in life: Dorsal surface with small granules; back with a dorsal ridge, in >-< shape; dorsolateral folds prominent; supratympanic fold distinct; flank with small tubercles; dorsal surface of limbs granular with short transverse ridges; throat, chest, belly and ventral surface of thigh smooth. Coloration in life: Dorsum reddish brown with a dark brown triangle between the eyes; a dark brown marking edged along dorsal ridge; flanks yellowish brown with some small black spots; tympanum dark brown; pupil black, outlined in reddish brown, iris copper-brown with tiny dark reticulations spreading from pupil; upper lip with a continuous white stripe, running from the nostril to shoulder; small white spots on flank and back of thigh; dorsal surface of fore and hind limbs reddish brown with dark crossbars; throat and chest dark brown, pectoral glands white; anterior part of belly white with brown pattern, posterior part of belly white; ventral surface of limbs dark brown with white marking, anterior part of thigh, posterior part of thigh dark brown; femoral glands white; toe webbing dark brown. Coloration in preservative: Dorsum greyish with a dark brown triangle marking between the eyes; a dark brown marking edged along dorsal ridge; flanks greyish with some small dark spots; throat and chest brown; belly cream with brown pattern; dorsal surface of fore and hind limbs greyish with dark brown crossbars; posterior part of thigh dark brown; femoral gland white; toe webbing dark brown. Variation. Measurements and morphological characters of the type series are given in Table 3. Adult males are smaller than adult females. Dorsum is reddish brown or grayish brown. Comparisons. We compared the new species with other species of Xenophrys from Vietnam and neighboring countries, including Laos, Cambodia, Thailand, and China. Xenophrys truongsonensis sp. nov. can be distinguished from its congeners by morphological characters as shown in Table 4. Morphologically, Xenophrys truongsonensis sp. nov. is most similar to X. maosonensis Bourret but they are different from each other by having flank with small tubercles (vs. flank with large tubercles in X. maosonensis), and toes with narrow lateral fringes (vs. absent in X. maosonensis), and white spots on flank and back of thigh smaller than those in X. maosonensis. Xenophrys truongsonensis sp. nov. differs from X. aceras Boulenger by having a greater TbL/SVL ratio in males (0.55–0.64 vs. 0.42–0.46 in X. aceras), and a greater ratio HL/HW>1 (vs. M. aceras); from X. ancrae Mahony, Teeling & Biju by having a larger body size (SVL 58.8–71.4 mm in males, 65.6–87.3 mm in females vs. 39.1–45.0 in males, 48.9 mm in female in X. ancrae), a greater ratio of TbL/SVL in males (0.55–0.64 vs. 0.48 in X. ancrae), toes with narrow lateral fringes (vs. absent in X. ancrae), and the absence of nuptial pads on finger I (vs. present in X. ancrae); from X. auralensis Ohler, Swan & Daltry by having a greater ratio of TbL/SVL in males (0.55–0.64 vs. 0.51 in X. auralensis), the presence of vomerine teeth (vs. absent in X. auralensis), the absence of external vocal sac (vs. present in X. auralensis), the first finger being shortest (vs. the second finger or fourth finger being shortest in X. auralensis), and tongue feebly notched (vs. tongue not notched in X. auralensis); from X. damrei Mahony by having a greater ratio of TbL/SVL in males (0.55–0.64 vs. 0.51 in X. damrei), the absence of external vocal sac (vs. present in X. damrei), and the first finger being shortest (vs. the fourth finger being shortest in X. damrei); from X. glandulosa Fei, Ye, & Huang by the absence of external vocal sac (vs. present in X. glandulosa), toes with narrow lateral fringes (vs. well developed fringes in X. glandulosa), and the first finger being the shortest (vs. the second finger or fourth finger being shortest in X. glandulosa); from X. lekaguli Stuart, Chuaynkern, Chanard, & Inger by having a greater ratio of TbL/SVL (0.55–0.64 in males and 0.54–0.58 mm in females vs. 0.47–0.52 in males and 0.44–0.52 in females in X. lekaguli), tongue feebly notched (vs. tongue not notched in X. lekaguli), toes with narrow lateral fringes (vs. absent in X. lekaguli), and the first finger being shortest (vs. the fourth finger being shortest in X. lekaguli); from X. longipes Boulenger by having a larger body size in males (SVL 58.8–71.4 mm vs. 47 mm in the male X. longipes), the absence of external vocal sac (vs. present in X. longipes), and the first finger being shortest (vs. the second finger being shortest in X. longipes); from X. mangshanensis Fei & Ye by the absence of external vocal sac (vs. present in X. mangshanensis), toes with narrow lateral fringes (vs. absent in X. mangshanensis), toes with rudiment of webbing (vs. without webbing in X. mangshanensis), and the fisrt finger being shortest (vs. the second finger being shortest in X. mangshanensis); from X. medogensis Fei, Ye, & Huang by the absence of external vocal sac (vs. present in X. medogensis), and toes with narrow lateral fringes (vs. absent in X. medogensis); from X. pachyproctus Huang by having a larger body size (SVL 58.8–71.4 mm in males and 65.6–87.3 mm in females vs. 35.3–36.2 mm in males and 35.8 mm in the female in X. pachyproctus), a greater ratio of TD/ED (0.51–0.67 in males and 0.53–0.59 in females vs. 0.35 in the male and 0.36 in the female in X. pachyproctus), a greater ratio of TbL/SVL (0.55–0.6 in males and 0.54–0.58 mm in females vs. 0.48–0.49 in males and 0.5 in the female in X. pachyproctus), the presence of vomerine teeth (vs. absent in X. pachyproctus), toes with narrow lateral fringes (vs. absent in X. pachyproctus), the absence of subarticular tubercles on toes (vs. present in X. pachyproctus), and the absence of nuptial pads on finger I (vs. present in X. pachyproctus); from X. takensis Mahony by having a larger body size in males (58.8–71.4 mm vs. 47.3–53.0 mm in X. takensis), a greater ratio of TD/ED in males (0.51– 0.67 vs 0.42–0.48 in X. takensis), a greater ratio of TbL/SVL (0.55–0.64 in males and 0.54–0.58 mm in females vs. 0.41–0.49 in males and 0.45 in female in X. takensis), the absence of external vocal sac (vs. present in X. takensis), tongue feebly notched (vs. tongue not notched in X. takensis), toes with narrow lateral fringes (vs. absent in X. takensis), the first finger being shortest (vs. the fourth finger being shortest in X. takensis), and the absence of nuptial pads on finger I (vs. present in X. takensis); from X. yeae by having a larger body size (58.8–71.4 mm in males and 65.6–87.3 mm in females vs. 23.0– 29.1 mm in males and 27.9–31.1 mm in the females in X. yeae), a greater ratio of TD/ED (0.55–0.64 in males and and 0.53–0.59 in females vs. 0.38–0.47 in males and 0.39–0.50 in females in X. yeae), and the presence of vomerine teeth (vs. absent in X. yeae); from X. zhangi Ye & Fei by having a larger body size in males (58.8–71.4 mm vs. 32.5–37.2 mm in X. zhangi), a greater ratio of TD/ED in males (0.51–0.67 vs 0.5 in X. zhangi), a greater ratio of TbL/SVL in males (0.55–0.64 vs. 0.49 in X. zhangi), the absence of external vocal sac (vs. present in X. zhangi), and the absence of nuptial pads on finger I (vs. present in X. zhangi); and from X. zhoui by having a larger body size (58.8–71.4 mm in males and 65.6–87.3 mm in females vs. 23 mm in male and 23.5–23.9 mm in the females in X. zhoui), a greater ratio of TD/ED in males (0.55–0.64 vs. 0.48 in X. zhoui), the presence of vomerine teeth (vs. absent in X. yeae), and toes with narrow lateral fringes (vs. absent in X. zhoui). ......continued on the next page ......continued on the next page Etymology. Specific epithet is in reference to the type locality of the Truong Son Mountain Range. We recommend “Truongson Horned Toad” as the common English name and “Cóc mắt trường sƠn” as the Vietnamese name. Ecological notes. Specimens were collected at night between 19:00 and 23:00 on tree leaves, ca 0.2–0.6 m above the ground, or on ground in rocky streams. The surrounding habitat was secondary evergreen forest consisting of medium and small hardwoods mixed with shrubs or secondary evergreen forest consisting of small hardwoods mixed with shrubs (Fig. 5). Distribution. Xenophrys truongsonensis sp. nov. is currently known from Thua Thien Hue Province southwards to Lam Dong Province, Central Vietnam (Fig. 6). It is expected that this species can be found in the evergreen forests on western side of Truong Son Range in Laos and Cambodia as well.

Theloderma khoii sp. nov. urn:lsid:zoobank.org:act: 491C280C-F0CA-475E-8DC9-377603DCA558 Figs 2–3 Diagnosis Theloderma khoii sp. nov. shows the diagnostic characters of the genus Theloderma, for instance a distinct tympanum, round canthus rostralis, bony ridges from canthus rostralis to occiput absent, and skin of head not co-ossified to the skull, and also molecular analyses revealed it to be nested within Theloderma. The new species is distinguished from its congeners and other small rhacophorid species by a combination of the following characters: 1) large-sized frog of the genus Theloderma (SVL 52.2 mm in a single male, 59.4 mm in a single female); 2) head length and width equal; 3) vomerine teeth present; 4) snout pointed and truncated (SNL/SVL 16.3% in male; 16.0% in female); 5) eye large, eye diameter about a half of snout length (ED/SNL 54.60% in male; 58.9% in female), spinules on upper eyelid; 6) tibiotarsal articulation reaching to the posterior border of the eye or the tip of the snout; 7) dorsal skin very rough, with large irregular gland ridges and warts, ventral surface granular; 8) fingers rudimentary webbed, toes almost four-fifths webbed, tips of all digits dilated but all considerably smaller than tympanum; 9) dorsal surface mossy green or olive mottled with dark magenta; 10) lateral and ventral surfaces black with irregular lemon yellow pattern and some white granules; 11) male with large nuptial pads and an inner vocal sac. Etymology The specific epithet is in honor of Professor Dr Le Vu Khoi of the VNU Hanoi University of Science, Ha Noi, Vietnam, in recognition of his great support of teaching, mentoring, research and conservation work in Vietnam. As common names we suggest Khoi’s Mossy Frog (English) and Ếch cây sần khôi (Vietnamese). Type material Holotype VIETNAM • adult &male;; northern Vietnam, Ha Giang Province, Quan Ba District, in the karst forest near Tung Vai Commune; 1641 m a.s.l; 8 Jun. 2020; H.Q. Nguyen leg.; VNMN 012757. Paratype VIETNAM • adult &female;; same collection data as for holotype; VNMN 012758. Description Holotype SIZE. Large, body distinctly flattened (SVL 52.2 mm). HEAD. Very strongly depressed, length and width equal (HL 19.7 mm, HW 19.7 mm), snout pointed and truncated, snout length double the diameter of eye (SNL 8.5 mm, ED 4.7 mm). CANTHUS ROSTRALIS. Oval, loreal region very oblique, slightly concave; interorbital distance wider than internarial distance and upper eyelid width (IOD 6.0 mm, IN 3.7 mm, UEW 4.5 mm); distance between anterior corners of eyes (DAE 10.1 mm) about 59.8% distance between posterior corners of eyes (DPE 16.3 mm). NOSTRIL. Oval, inside rather enlarged tubercle on tip of snout, nostrils nearer to tip of snout than to eyes (NS 3.1 mm, EN 6.1 mm). PUPIL. Circular. TYMPANUM. Distinct, diameter less than eye diameter and slightly larger than width of third finger disc (TYD 3.7 mm), tympanum separated from eye by distance (TYE) 65.0% of tympanum length (TYD). PINEAL OCELLUS. Absent; conical tubercles on eyelid, smaller than those in the back. SUPRATYMPANIC FOLD. Distinct, interrupted, extending from behind eye to beyond level of axilla, composed of large irregular glandular ridges. VOMERINE TEETH. Arranged in two small oblique groups between choanae and widely separated from each other. TONGUE. Large, accounting for almost half of mouth, heart-shaped, dorsal surface smooth, notched and free posterior. FORELIMBS. Moderately strong and long (FLL 13.2 mm, HAL 15.1 mm, 54.3% of SVL), much shorter than hindlimbs (FLL 87.5% of HAL); relative length of fingers: I large, rounded and flattened discs, with grooves separating dorsum of discs from venter; disc of finger III approximately two times width of finger III (fd3/fw3 219.2%) but smaller than tympanum diameter (fd3/TYD 83.1%); subarticular tubercles large and distinct, with indistinct supernumerary tubercles; formula of subarticular tubercles: 1, 1, 2, 2; well-developed nuptial pad on lateral side of thumb; outer side of fourth finger fringed, outer edge of fourth finger distinctly serrated; webbing between fingers slightly developed, about 50% between base of finger up to proximal subarticular tubercles (Fig. 3). HINDLIMBS. Strong and long, tibia length about five times greater than tibia width (TbL 27.3 mm, TbW 5.65 mm, TbL/TbW 4.6), tibia longer than thigh length and foot length (TbL/FeL 107%; TbL/ FoL 108%), tibio-tarsal articulation reaching posterior edge of snout when hindlimbs pressed forward as opposed to eye; relative length of toes I SKIN TEXTURE IN LIFE. Dorsal surface of head, dorsum, arms and legs above very rough, with large irregular glandular ridges ordered symmetrically in middle vertebral region; a large, distinct horn gland behind head in X-shape; on top of each supraorbital 3–4 enlarged glands, distinctly conical; lateral parts of body become granular; large conical tubercles on back of thighs near vent; throat and chest with some small blurred pattern; belly and ventral surface and underside of thigh with thickened flat granules or warts on groin; axillary region and underside of tibia smooth; surroundings of tympanum with small tubercles, with 4–5 enlarged tubercles posteriorly. COLORATION IN LIFE. Dorsal surface mossy-green or light-olive mixed with mossy-green, resembling moss; tips of skin ridges and tubercles reddish-brown; flank with clearly shaped dark brown pattern, ventrally joining a yellowish-green band; dorsal surfaces of forelimbs and hindlimbs with army-green bands, and some irregular patches on webbing; both finger and toe discs green, in center deep olive; nuptial pad ivory; throat and chest dark-brown with yellowish-green warts; belly has black shape of irregular size with yellowish-green pattern and some cream warts; pupil black, iris yellowish-green with irregular black reticulation; tympanum dark-olive (Fig. 2). COLORATION IN PRESERVATIVE. In ethanol, dorsal surfaces of head, dorsum, and upper part of flanks dark indigo with brown markings and blotches; tympanum light brown; forelimb, dorsal surface of thigh, tibia and foot black with lights bands, posterior part of thigh below vent dark-grey with large ivory bands; chin and throat light-grey with small ivory spots; chest and belly dark to black, with cream areas on chest (Fig. 3). Variation of paratype Male smaller than female, with single vocal sac and nuptial pads. For measurements of the type series see Table 3. Comparison with other species We compared Theloderma khoii sp. nov. with other members of the genus Theloderma based on morphological specimen examination (see Appendix 1) and data obtained from the literature (e.g., Tschudi 1838; Boulenger 1903; Smith 1924; Ahl 1927, 1931; Bourret 1937, 1942; Liu & Hu 1962; Taylor 1962; Chanda 1994; Inger et al. 1999; Chanard 2003; Bain & Nguyen 2004; Stuart & Heatwole 2004; Orlov & Ho 2005; Orlov et al. 2006, 2012; McLeod & Norhayati 2007; Bain et al. 2009; Chan & Norhayati 2009; Fei et al. 2009, 2012; Jiang et al. 2009; Kunz et al. 2010; Rowley et al. 2011; Nguyen et al. 2014). Species of Theloderma inhabiting Indochina, Sumatra, southern China and eastern India can be assigned to three different size groups (Dever 2017): small species (SVLT. albopunctatum (Liu & Hu, 1962); T. annae Nguyen, Pham, Nguyen, Ngo & Ziegler, 2016; T. asperum (Boulenger, 1886); T. auratum Poyarkov, Kropachev, Gogoleva & Orlov, 2018; T. baibungense Jiang, Fei & Huang, 2009; T. lacustrinum Sivongxay, Davankham, Phimmachak, Phoumixay & Stuart, 2016; T. laeve (Smith, 1924); T. lateriticum Bain, Nguyen & Doan, 2009; T. licin McLeod & Ahmad, 2007; T. nebulosum and T. palliatum Rowley, Le, Hoang, Dau & Cao, 2011; T. petilum (Stuart & Heatwole, 2004); T. pyaukkya Dever, 2017; T. rhododiscus (Liu & Hu, 1926); T. (Stelladerma in Poyarkov et al. 2015) stellatum Taylor, 1926; T. (Stelladerma in Poyarkov et al. 2015) vietnamense (Poyarkov et al., 2015); and T. truongsonense (Orlov & Ho, 2005); medium-sized species with SVL of 40–45 mm, T. horridum (Boulenger, 1903); T. moloch (Annandale, 1912); T. phrynoderma (Ahl, 1927); T. nagalandense and T. ryabovi Orlov, Dutta, Ghate & Kent, 2006; and T. schmardana (Kelaart, 1853); large-sized species with SVL 48–75 mm (Orlov 1997), including T. bicolor (Bourret, 1937); T. corticale (Boulenger, 1903); T. gordoni Taylor, 1962; T. leporosum Tschudi, 1838; and Theloderma khoii sp. nov. Compared to T. bicolor, another large-sized group member in Vietnam, Theloderma khoii sp. nov. can be distinguished by its coloration pattern: the lateral part from the axilla to groin is black with an irregular yellowish-green pattern in T. bicolor vs flank pattern with clear brown-dark speckles, ventrally joining the yellowish-green band in Theloderma khoii sp. nov.; a comparatively lighter belly pattern, consisting of irregular blackish brown speckles with whitish pattern in T. bicolor vs belly irregular black with yellowish-green pattern and some cream warts, and the presence of few smaller bluish spots on the throat, in Theloderma khoii sp. nov. (Fig. 4); by the comparatively dark-colored posterior part of the thighs, consisting of black blotches and irregular light-green or olive reticulations (vs black with yellowgreenish bands and a few red-brown warts in Theloderma khoii sp. nov.; Fig. 4); and foot webbing well-developed: I(0) – (1)II(0) – (1/2)III(0) – (1)IV(1) – (0)V (vs I(1/3) – (1)II(0) – (1/2)III(0) – (1) IV(1) – (1/3)V in Theloderma khoii sp. nov.; Fig. 4). Theloderma khoii sp. nov. (Bourret 1937; Hou et al. 2017) differs from T. corticale by its shorter body size: SVL 52.2 mm in male and 59.4 mm in female in Theloderma khoii sp. nov. vs SVL up to 67 mm in T. corticale; by colour pattern: a much lighter belly pattern, yellow-lemon with irregular small black patterns and creamy warts (vs dark belly pattern, blackish ground color with thin yellowish-green pattern and some cream warts in Theloderma khoii sp. nov.); by skin texture: the absence of enlarged warts in the scapular area (vs large irregular prominent warts on dorsum, forming large groups at the base of the head and scapular area in Theloderma khoii sp. nov.); by the head being slightly wider than long (vs head length and width equal in Theloderma khoii sp. nov.); and foot webbing well developed: webbing formula: I(0) – (1 + 12)II(0) – (1)III(0) – (1) IV(1) – (0)V (vs I(1/3) – (1)II(0) – (1/2)III(0) – (1)IV(1) – (1/3)V in Theloderma khoii sp. nov. Theloderma khoii sp. nov differs from T. gordoni by its coloration pattern: dorsal surface mossy-green or light-olive mixed with mossy-green, resembling moss; ventral yellowish-green band in Theloderma khoii sp. nov. vs dorsum dark brown or coffee with some clusters light brown or orange on enlarged gland ridges; ventral surface dark blue with numerous irregular grayish white patterns and speckles in T. gordoni. Tympanum and tympanic fold distinct in Theloderma khoii sp. nov. vs tympanum and tympanic fold invisible in T. gordoni; head width equal to head length in Theloderma khoii sp. nov. vs head width greater than head length in T. gordoni (Qui et al. 2018). Ecological notes Theloderma khoii sp. nov. appears to be closely associated with karstic environments. Specimens were found at night between 19:00 and 23:30 h near cave entrances and in valleys surrounded by limestone cliffs, about 5–6 m from water sources (Fig. 5). Advertisement calls, eggs and tadpoles of the species have not been recorded during our field surveys. The main habitat at the type locality was secondary karst forest of medium and small hardwoods mixed with shrubs and vines. One specimen was found on a leaf, about 0.5–1.2 m above the ground, the other specimen was collected on a limestone cliff, about 0.5–1.2 m above the ground. The air temperature at the time of collection ranged from 18.3 to 23.2 oC and relative humidity from 91 to 100%. In Tung Vai, several species of Theloderma have been recorded, viz. T. albopunctatum (Liu & Hu, 1962), T. gordoni (Taylor, 1962), T. lateriticum (Bain, Nguyen & Doan, 2009), and T. rhododiscus (Liu & Hu, 1962), but all were found in tree holes filled with water or on tree leaves near streams in the valleys. Other tree frogs that were found at the same site were Polypedates sp. (of the P. megacephalus Hallowell, 1861 species complex), Zhangixalus duboisi (Ohler, Marquis, Swan & Grosjean, 2000), Z. jodiae Nguyen, Ninh, Orlov, Nguyen & Ziegler, 2020, Z. franki Ninh, Nguyen, Orlov, Nguyen & Ziegler, 2020, Z. pachyproctus Yu, Hui, Hou, Wu, Rao & Yang, 2019 and Kurixalus sp. Distribution Theloderma khoii sp. nov. is currently known only from the type locality in Ha Giang Province, northeastern Vietnam (Fig. 6). The species was recorded at elevations between 1320 and 1750 m a.s.l. Conservation status The range of the new species is not expected to extend outside of Mount Tung Vai and it is expected to be found in evergreen forests of Yunnan Province, southern China. However, the actual distributional range should be the focus of further studies. Currently, we assume that the new species is restricted geographically, likely having an Extent Of Occurrence (EOO) of only et al. 2013). Therefore, Theloderma khoii sp. nov. likely qualifies as Endangered (EN) B1ab(iii) in accordance with the categories and criteria of the IUCN Red List of Threatened Species (IUCN 2012).

Lötters, Stefan, Plewnia, Amadeus, Hönig, Alana, Jung, Alisha, Laudor, Jörn, Ziegler, Thomas (2022): -The-gastromyzophorous-tadpole-of-the-pink-harlequin-frog-from-Suriname-withcomments-on-the-taxonomy-of-Guianan-Clade-Atelopus-(Amphibia,-Bufonidae). Zootaxa 5087 (4): 591-598, DOI: https://doi.org/10.11646/zootaxa.5087.4.7

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"]}

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; Rösler 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 (rösler 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 Rösler, 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 Günther, 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.

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 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.

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"]}

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 Thằn lằn ngón ngật (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.

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.

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., Published as part of Nguyen, Thuong Huyen, Luu, Vinh Quang, Sitthivong, Saly, Ngo, Hanh Thi, Nguyen, Truong Quang, Le, Minh Duc & Ziegler, Thomas, 2021, A new species of Dixonius (Squamata: Gekkonidae) from Vientiane Capital, Laos, pp. 351-362 in Zootaxa 4965 (2) on pages 354-360, DOI: 10.11646/zootaxa.4965.2.8, http://zenodo.org/record/4750274

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 • &male;; 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 &female;; 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).

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 Th���ch s��ng ng��n ch���ng (Vietnamese). Type material (Figs 3���8) Holotype VIETNAM ��� &male;; 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 &female;; 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; R��sler 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, R��sler, Vu, B��hme & Ziegler, 2007, C. gialaiensis Luu, Tran, Nguyen, Le & Ziegler, 2017, C. pseudoquadrivirgatus R��sler, 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)., Published as part of 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, pp. 47-70 in European Journal of Taxonomy 731 on pages 53-64, DOI: 10.5852/ejt.2021.731.1203, http://zenodo.org/record/4438108, {"references":["Smith M. A. 1921 a. New or little-known reptiles and batrachians from Southern Annam (Indo-China). Proceedings of the Zoological Society of London 91 (2): 423 - 440. https: // doi. org / 10.1111 / j. 1096 - 3642.1921. tb 03271. x","Smith M. A. 1921 b. Reptiles and batrachians collected on Pulo Condore. The Journal of the Natural History Society of Siam 4 [1920]: 93 - 97. Available from https: // www. biodiversitylibrary. org / part / 75704 / summary [accessed 21 Nov. 2020].","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., 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","Heidrich A., Rosler H., Vu T. N., Bohme W. & Ziegler T. 2007. Another new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha - Ke Bang National Park, central Truong Son, Vietnam. Zootaxa 1445 (1): 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1445.1.3","Orlov N. L., Nguyen Q. T., 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 (2): 145 - 152.","Nazarov R. A., Orlov N. L., Nguyen N. S. & Ho T. C. 2008. Taxonomy of naked-toes 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 (2): 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 (1): 27 - 42. https: // doi. org / 10.11646 / zootaxa. 1715.1.2","Rosler H., Ngoc T. V., Nguyen T. Q., Ngo V. T. & Ziegler T. 2008. A new Cyrtodactylus (Squamata: Gekkonidae) from Central Vietnam. Hamadryad 33 (1): 48 - 63.","Geissler P., Nazarov R., Orlov N., Bohme W., Phung T. M., Nguyen T. Q. & Ziegler T. 2009. A new species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from southern Vietnam. Zootaxa 2161 (1): 20 - 32. https: // doi. org / 10.11646 / zootaxa. 2161.1.2","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. 2013. Cyrtodactylus dati, a new forest dwelling bent-toed gecko (Squamata: Gekkonidae) from southern Vietnam. Zootaxa 3616 (2): 151 - 164. https: // doi. org / 10.11646 / zootaxa. 3616.2.4","Nguyen S. N., Le T. T., Tran T. A. D., Orlov N. L., Lathrop A., Macculloch R. D., Le T. T., Jin J., Nguyen L. T., Nguyen T. T., Hoang D. D., Che J., Murphy R. W. & Zhang Y. - P. 2013. Phylogeny of the Cyrtodactylus irregularis species complex (Squamata: Gekkonidae) from Vietnam with the description of two new species. Zootaxa 3737 (4): 399 - 414. https: // doi. org / 10.11646 / zootaxa. 3737.4.4","Schneider N., Phung T. M., Le M. D., Nguyen T. Q. & Ziegler T. 2014. A new Cyrtodactylus (Squamata: Gekkonidae) from Khanh Hoa Province, southern Vietnam. Zootaxa 3785 (4): 518 - 532. https: // doi. org / 10.11646 / zootaxa. 3785.4.2","Luu V. Q., Tran D. 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","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","Neang T., Henson A. & Stuart B. L. 2020. A new species of Cyrtodactylus (Squamata, Gekkonidae) from Cambodia's Prey Lang Wildlife Sanctuary. ZooKeys 926: 133 - 158. https: // doi. org / 10.3897 / zookeys. 926.48671","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","Ngo T. V. 2008 a. Two new cave-dwelling species of Cyrtodactylus Gray (Squamata: Gekkonidae) from Southwestern Vietnam. Zootaxa 1909 (1): 37 - 51. https: // doi. org / 10.11646 / zootaxa. 1909.1.4","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 (3): 215 - 226.","Nguyen S. N., Yang J-X., Le N. T. 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 benttoed 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., TranA. - 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. Cytonuclear discordance, cryptic diverity, complex histories and conversation needs on vietnamese bent-toed geckos of the Cyrtodactylus irregularis complex. Russian Journal of Herpetology 24 (2): 133 - 154.","Ngo T. V. 2008 b. Cyrtodactylus martini, another new karst-dwelling Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Northwestern Vietnam. Zootaxa 2834 (1): 33 - 46. https: // doi. org / 10.11646 / zootaxa. 2834.1.3","Ngo H. T., Le M. D., Pham N. D., Nguyen V. T. H., Pham C. T. & Nguyen T. Q. 2017. Taxonomy and phylogenetic relationships of Cyrtodactylus (Squamata: Gekkonidae) in Vietnam. VNU Journal of Science: Natural Sciences and Technology 33 (1 S): 182 - 192. https: // doi. org / 10.25073 / 2588 - 1140 / vnunst. 4530","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","Ngo T. V. & Grismer L. L. 2012. A new endemic 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, Vietnam. Zootaxa 2413 (1): 20 - 36. https: // doi. org / 10.11646 / zootaxa. 2413.1.2","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 (1): 39 - 50. https: // doi. org / 10.11646 / zootaxa. 3129.1.3","Ngo T. V. & Chan K. O. 2011. A new karstic cave-dwelling Cyrtodactylus Gray (Squamata: Gekkonidae) from Northern Vietnam. Zootaxa 3125 (1): 51 - 63. https: // doi. org / 10.11646 / zootaxa. 3125.1.4","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. http: // dx. doi. org / 10.11646 / zootaxa. 3764.3.5","Grismer L. L., Wood P. L. Jr., 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","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 east-central and southern Myanmar demonstrate high localized diversity and unprecedented microendemism. Zoological Journal of the Linnean Society 182 (4): 862 - 959. https: // doi. org / 10.1093 / zoolinnean / zlx 057","Nguyen T. Q., Le M. D., Pham A. V., Ngo H. N., Hoang C. V., Pham C. T. & Ziegler T. 2015. Two new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of Hoa Binh Province, Vietnam. Zootaxa 3985 (3): 375 - 390. https: // doi. org. / zootaxa. 3985.3.3","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","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","Murdoch M. L., Grismer L. L., Wood P. L. Jr., Neang T., Poyarkov N. A., Ngo V. T., 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"]}

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

Ziegler, Thomas, Pham, Cuong The, Nguyen, Tan Van, Nguyen, Truong Quang, Wang, Jian, Wang, Ying-Yong, Stuart, Bryan L., Le, Minh Duc (2021): THOMAS ZIEGLER, CUONG THE PHAM, TAN VAN NGUYEN, TRUONG QUANG NGUYEN JIAN WANG, YING-YONG WANG, BRYAN L. STUART & MINH DUC LE (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, 579-586. Zootaxa 4903 (4): 598-598, DOI: https://doi.org/10.11646/zootaxa.4903.4.8

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).

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)., Published as part of 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, pp. 413-422 in Revue suisse de Zoologie 127 (2) on pages 416-420, DOI: 10.35929/RSZ.0030, http://zenodo.org/record/5743934, {"references":["Janssen H. Y., Pham C. T., Ngo H. T., Le M. D., Nguyen T. Q., Ziegler T. 2019. A new species of Lycodon Boie, 1826 (Serpentes, Colubridae) from northern Vietnam. Zookeys 875: 1 - 29.","Ye C., Fei L., Hu S. 1993. Rare and economic amphibians of China. Sichuan Publishing House of Science & Technology, Chengdu, 412 pp. [in Chinese]","Ohler A., Delorme M. 2006. Well known does not mean well studied: Morphological and molecular support for existence of sibling species in the Javanese gliding frog Rhacophorus reinwardtii (Amphibia, Anura). Comptes Rendus Biologies 329: 86 - 97.","Gray J. E. 1845. Catalogue of the specimens of lizards in the collection of the British Museum. Trustees of the British Museum / Edward Newman, London: xxvii + 289 pp.","Cuvier G. L. 1829. Le Regne Animal Distribue, d'apres son Organisation, pour servir de base a l'Histoire naturelle des Animaux et d'introduction a l'Anatomie Comparee. Nouvelle Edition [second edition]. Vol. 2. Les Reptiles. Deterville, Paris, i-xvi, 1 - 406.","Grismer 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.","Boie F. 1827. Bemerkungen uber Merrem's Versuch eines Systems der Amphibien, 1. Lieferung: Ophidier. Isis van Oken 20: 508 - 566."]}

Limnonectes phuyenensis sp. nov. (Figs. 2–5) Holotype: IEBR 4737 (Field number DTD 138), adult male, collected by D. T. Do on 28 March 2015 (12°51.184 ’ N, 109°13.151 ’ E, at an elevation of 359 m asl.) in the evergreen forest of Ca Mountain Pass, near Hoa Thinh Commune, Tay Hoa District, Phu Yen Province, Vietnam. Paratypes: 18 specimens collected from the evergreen forest of Phu Yen Province, Vietnam: One adult male, PYU 535 (Field number DTD 04) and two adult females, IEBR 4739, 4740 (Field numbers DTD 163, 171), collected by D. T. Do on 28 March 2015 (12°51.184 ’ N, 109°13.151 ’ E, at an elevation of 359 m asl.) from Ca Mountain Pass, near Hoa Thinh Commune, Tay Hoa District; one adult male, IEBR 4738 (Field number DTD 535) and one adult female, PYU 536 (Field number DTD 536), collected by D. T. Do on 19 August 2015 (12°52.470 ’ N, 108°49.511 ’ E, at an elevation of 554 m asl.) from Ca Mountain Pass, near Son Thanh Tay Commune, Tay Hoa District; one adult male, ZFMK 103169 (Field number DTD 680) and one adult female, ZFMK 103170 (Field number DTD 708), collected by D. T. Do on 10 September 2016 (13°27.437 ’ N, 108°52.005 ’ E, at an elevation of 430 m asl.) from Phu Mo Commune, Dong Xuan District; one adult male, ITBCZ 4598 (Field number SH 040) collected by L. T. Nguyen, M. V. Le, and H. V. Lo on 26 March 2019 (12°78.549 ’ N, 109°03.737 ’ E, at an elevation of 350 m asl.); six adult males, ITBCZ 4746, 4747, 4749, 4753, 4754 (Field numbers SH 188, 189, 191, 195, 196, respectively) and VNMN 010902 (Field number SH194) and four adult females, ITBCZ 4745, 4748, 4750 (Field numbers SH 187, 190, 192, 193, respectively) and VNMN 010903 (Field number SH193) collected by L. T. Nguyen, M. V. Le, and H. V. Lo on 31 March 2019 (12°78.549 ’ N, 109°03.737 ’ E, at an elevation of 350 m asl.) from Song Hinh Protection Forest, Song Hinh District. Diagnosis: Molecular and morphological data, including the presence of fang-like odontoid processes on the lower jaw and sexual dimorphism, with males larger than females and enlargement of the head in males (Emerson et al. 2000), revealed the new species to be nested within Limnonectes. Limnonectes phuyenensis sp. nov. is distinguishable from its congeners by a combination of the following morphological characters: (1) SVL 49.8–77.9 mm in males, 40.8–61.8 mm in females; (2) males with moderately enlarged head (HL/SVL 0.47, HW/SVL 0.48); (3) head slightly broader than long (HL/HW 0.97 in males, 0.95 in females); (4) vomerine teeth present; (5) external vocal sacs absent; (6) rostral length short (RL/SVL 0.15); (7) tympanum invisible; (8) dorsal surface of head, body and flanks with ridges and tubercles; (9) dorsal surface of tibia and foot distinctly tuberculate, covered with moderately dense, small, low tubercles; (10) supratympanic fold present; (11) dorsolateral fold absent; (12) Finger I with nuptial pad, composed of minute spines on dorsal surface and medial edge in males; (13) toes webbed to distal of terminal phalanx, webbing formala I0–0II0–0III0–0IV0 - –0V0; (14) in life, dorsum yellowish brown with dark brown markings; (15) ventral surface white with brown markings. Description of holotype: Adult male (IEBR 4737); SVL 62.8 mm; habitus robust with enlarged head (HL/SVL 0.48, HW/SVL 0.50); head slightly broader than long (HL 30.1 mm, HW 31.2 mm); snout round anteriorly in dorsal view, projecting beyond lower jaw; nostril lateral, closer to tip of snout than to eye (NS 4.2 mm, EN 4.8 mm); canthus rostralis indistinct; loreal region oblique and slightly concave; rostral length greater than eye diameter (RL 9.0 mm, ED 7.6 mm); interorbital distance wider than internarial distance and upper eyelid width (IOD 6.4 mm, IND 5.3 mm, UEW 3.9 mm); tympanum invisible; vomerine teeth in two oblique ridges; tongue cordiform, notched posteriorly; lower jaw with two tooth-like, odontoid processes robust with very sharp tips (OH 4.0 mm); external vocal sac absent. Forelimbs: Arms short; upper arm length (UAL) 10.8 mm, forearm length (FAL) 26.2 mm; relative finger lengths: I Hind limbs: Tibia length shorter than thigh length (FeL 28.8 mm, TbL 27.1 mm), approximately 2.2 times longer than wide (TbW 12.4 mm); tips of toes blunt, slightly round; relative length of toes: I Skin texture in life: Dorsal surface of head, body and flanks with ridges and tubercles; small tubercles on upper eyelid; supratympanic fold distinct, extending from eye to angle of jaw; dorsolateral fold absent; small tubercles around cloaca; dorsal surface of forelimbs and thighs smooth; dorsal surface of tibia and foot distinctly tuberculate, covered with moderately dense, small, low tubercles; throat, chest, belly and ventral surface of thighs smooth. Coloration in life: Iris dark gray in upper half and light gray in lower half; head with a pale yellow bar in anterior interorbital region and a narrow dark brown bar in posterior interorbital region; dorsum and flanks with yellowish brown marking; lips with dark bars; supratympanic fold dark brown; dorsal surface of limbs yellowish brown with dark crossbars; ventral surface of limbs, throat, chest, and belly white with brown markings; toe webbing brown. Coloration in preservative: Dorsum and upper part of flanks pale brown with dark brown marking; lower part of flanks white with brown marking; lips with dark bars; head with a narrow dark brown bar in anterior interorbital region; supratympanic fold dark brown; dorsal surface of limbs pale brown with dark crossbars; ventral surface of limbs, throat, chest, and belly white with brown markings; minute spines on Finger I greyish brown; toe webbing brown. Variation and sexual dimorphism: Measurements and morphological characters of the type series are provided in Table 1 and photographs of paratypes in life are presented in Fig. 6. Males are larger than females (SVL 60.1 ± 8.3 mm, n = 11 males vs. 50.3± 7.3 mm, n = 8 females). Males have a larger head than females (HL/ SVL 0.47 vs. 0.44, HW / SVL 0.48 vs. 0.46). The male specimens have a nuptial pad on Finger I. The males have robust fanglike odontoid processes, which are less pronounced in the females. The females contained yellowish cream eggs of varying sizes. One juvenile (unvouchered) had a yellowish dorsum with distinct brown marking (Figure 6F). ......continued on the next page Etymology. The specific epithet “ phuyenensis ” refers to the type locality of the new species, Phu Yen Province. As common names we suggest Phu Yen Wart Frog (English), Ếch nhẽo phú yên (Vietnamese). Ecological notes. Specimens were found between 19:00 and 23:00 in the head-water of small, rocky streams (Fig. 7A). They were found in the water or on the ground of stream banks at elevations above 300 m. Males were sometimes found in small water pools with two or three females nearby. The surrounding habitat was secondary forest of large, medium and small hardwoods mixed with shrubs and vines. Air temperature at the sites was 20.5–26.3 oC and relative humidity was 73–96%. The females collected in March, August, and September contained yellowish cream eggs of varying sizes. Male advertisement calls and tadpoles of the species have not been recorded during our field surveys. Other amphibian species found at the sites included Leptobrachella macrops (Duong, Do, Ngo, Nguyen & Poyarkov), Megophrys elfina (Poyarkov, Duong, Orlov, Gogoleva, Vassilieva, Nguyen, Nguyen, Nguyen, Che & Mahony), Limnonectes poilani (Bourret), Odorrana morafkai (Bain, Lathrop, Murphy, Orlov & Ho), Kurixalus cf. banaensis (Bourret), and Rhacophorus annamensis (Smith). Distribution. Limnonectes phuyenensis sp. nov. is currently known only from Phu Yen Province, South-central Vietnam (Fig. 8). Comparisons. We compared the new species with its congeners from Vietnam and neighboring countries including Laos, Cambodia, Thailand, and China. Limnonectes phuyenensis sp. nov. differs from L. bannaensis Ye, Fei & Jiang by having a smaller size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 74.9 mm, n= 10 males, 60.8 mm, n= 8 females in L. bannaensis), head broader than long (vs. head longer than wide in L. bannaensis), tips of toes blunt (vs. tips of toes expanded into discs in L. bannaensis), dorsal surface of tibia and foot with tubercles (vs. smooth in L. bannaensis), and males with nuptial pad on Finger I (vs. males with nuptial pad on fingers I and II in L. bannaensis); from L. coffeatus Phimmachak, Sivongxay, Seateum, Yodthong, Rujirawan, Neang, Aowphol & Stuart by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 37.9 mm, n= 1 male, 40.5 mm, n= 3 females in L. coffeatus), males with nuptial pad on Finger I (vs. absence in males of L. coffeatus), tympanum invisible (vs. visible in L. coffeatus), and the absence of W marking on head (vs. presence in L. coffeatus); from L. dabanus (Smith) by the absence of caruncle on head in males (vs. presence in L. dabanus), tympanum invisible (vs. visible in L. dabanus), toes fully webbed (vs. webbing formula I0–1/3II0–1/4III0–1IV1–0V in L. dabanus), and males with nuptial pad on Finger I (vs. absence in L. dabanus); from L. fastigatus by having a differrent odontoid processes shape (robust with very sharp tips vs. thin and elongated in L. fastigatus), nostril closer to tip of snout than to eye (vs. midway between snout tip and eye in L. fastigatus), and belly with brown flecking or vermiform markings in preservative (vs. creamy white in preservative in L. fastigatus); from L. fragilis (Liu & Hu) by having a head broader than long (vs. head longer than wide in L. fragilis), a greater ratio of HL/SVL and HW/SVL (HL/SVL=0.47, HW/SVL=0.48, n= 11 males and HL/SVL=0.44, HW/SVL=0.46, n= 8 females vs. HL/SVL=0.45, HW/SVL=0.42, n= 6 males and HL/SVL=0.42, HW/SVL=0.41, n= 10 females in L. fragilis), tibia length shorter than thigh length (vs. tibia length longer than thigh length in L. fragilis), and males with nuptial pad on Finger I (vs. absence in males of L. fragilis); from L. fujianensis Ye & Fei by having a head broader than long (vs. head longer than wide in L. fujianensis), a greater ratio of HL/SVL and HW/SVL (HL/SVL=0.47, HW/SVL=0.48, n= 11 males and HL/SVL=0.44, HW/SVL=0.46, n= 8 females vs. HL/SVL=0.45, HW/SVL=0.44, n= 20 males and HL/SVL=0.38, HW/SVL=0.37, n= 20 females in L fujianensis), males with nuptial pad on Finger I (vs. nuptial pad on Fingers I and II in L. fujianensis), and the absence of W marking on head (vs. presence in L. fujianensis); from L. gyldenstolpei (Andersson) by the absence of caruncle on head in males (vs. presence in L. gyldenstolpei), tympanum invisible (vs. visible in L. gyldenstolpei), toes fully webbed (vs. webbing formula I0–3/4II0–1III1/4–1IV11/2–0V in L. gyldenstolpei), and males with nuptial pad on Finger I (vs. absence in L. gyldenstolpei); from L. kiziriani by having larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 54.1 mm, n= 13 males, 46.8 mm, n= 7 females in L. kiziriani), head broader than long (vs. longer than wide in L. kiziriani), a smaller ratio of TbL/TbW (TbL/TbW 2.31, n= 11 males and 2.40, n= 8 females vs. TbL/TbW 2.54, n= 13 males and 2.57, n= 7 females in L. kiziriani), and the absence of two dermal ridges parallel from posterior of eye to the end of shoulder (vs. presence in L. kiziriani); from L. hascheanus (Stoliczka) by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm n= 8 females vs. 22.1 mm, n= 9 males, 23.0 mm, n= 9 females of L. hascheanus), the absence of dorsolateral folds (vs. presence in L. hascheanus), males with nuptial pad on Finger I (vs. absence in males of L. hascheanus), and tympanum invisible (vs. visible in L. hascheanus); from L. isanensis McLeod, Kelly & Barley by having a smaller size in males (SVL 60.1 mm, n=11 vs. 71.5 mm, n= 11 in L. isanensis), head broader than long (vs. head longer than wide in L. isanensis), dorsum and flank with tubercles (vs. absence in L. isanensis), and males with nuptial pad on Finger I (vs. nuptial pad on Fingers I and II in L. isanensis); from L. jarujini Matsui, Panha, Khonsue & Kuraishi by having a smaller size in males (SVL 60.1 mm, n=11 vs. 73.3 mm, n= 8 in L. jarujini), head broader than long (vs. head longer than wide in L. jarujini), length of fingers IL. jarujini), and dorsum and flank with tubercles (vs. absence in L. jarujini); from L. khammonensis (Smith) by having a larger size in females (SVL 50.3 mm, n=8 vs. 37 mm, n= 1 in L. khammonensis) and tympanum invisible (vs. visible in L. khammonensis); from L. kohchangae (Smith) by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 35.9 mm, n= 21 males, 35.2 mm, n= 14 females of L. kohchangae), tympanum invisible (vs. visible in L. kohchangae) and males with nuptial pad on Finger I (vs. absent in L. kohchangae); from L. limborgi (Sclater) by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 33.0 mm, n= 30 males, 31.7 mm, n= 24 females of L. limborgi), the absence of dorsolateral folds (vs. presence in L. limborgi), males with nuptial pad on Finger I (vs. absence in males of L. limborgi), and tympanum invisible (vs. visible in L. limborgi); from L. liui (Yang) by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 35.7 mm, n= 3 males, 32.7 mm, n= 1 female of L. liui), tympanum invisible (vs. visible in L. liui), and dorsolateral fold absent (vs. present in L. liui); from L. longchuanensis Suwannapoom, Yuan, Sullivan & McLeod by having head broader than long (vs. head longer than wide in L. longchuanensis), males with nuptial pad on Finger I (vs. nuptial pad on Fingers I and II in L. longchuanensis), and dorsal ridges present (vs. absent in L. longchuanensis); from L. lauhachindai Aowphol, Rujirawan, Taksinum, Chuaynkern & Stuart by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 33.8 mm, n= 20 males, 35.2 mm, n= 4 females in L. lauhachindai), the absence of caruncle on head (vs. presence in L. lauhachindai), tympanum invisible (vs. visible in L. lauhachindai), and males with nuptial pad on Finger I (vs. absence in L. lauhachindai); from L. macrognathus (Boulenger) by having a larger size in males (SVL 60.1 mm, n=11 vs. 45.5 mm, n= 3 in L. macrognathus), the absence of caruncle on head (vs. presence in L. macrognathus), and tympanum invisible (vs. visible in L. macrognathus); from L. megastomias McLeod by having a smaller size (SVL 60.1 mm, n= 11 males, 50.3 mm in, n= 8 females vs. 80.2 mm, n= 22 males, 74.0 mm, n= 11 females), head broader than long (vs. head longer than wide in L. megastomias), dorsal surface of tibia and foot with tubercles (vs. smooth in L. megastomias), and males with nuptial pad on Finger I (vs. nuptial pad on Fingers I and II in L. megastomias); from L. nguyenorum by having a smaller ratio of HL/SVL (0.47, n= 11 males, 0.44, n= 8 females vs. 0.42–0.43, n= 2 males, 0.39–0.39, n= 2 females in L. nguyenorum), the head broader than long (vs. head longer than wide in L. nguyenorum), tibia shorter than thigh (vs. tibia longer than thigh in L. nguyenorum), males with nuptial pad on Finger I (vs. nuptial pad on Fingers I and II in L. nguyenorum), and the absence of two dermal ridges parallel from posterior of eye to the end of shoulder (vs. presence in L. nguyenorum); from L. poilani (Bourret) by having a smaller size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 90.0 mm, n= 4 males, 80.2 mm, n= 6 females of L. poilani), head broader than long (vs. head longer than wide in L. poilani), tibia shorter than thigh (vs. tibia longer than thigh in L. poilani), tympanum invisible (vs. visible in L. poilani), and ventral surface of limbs and abdomen white with brown markings (vs. yellow in L. poilani); from L. quangninhensis Pham, Le, Nguyen, Ziegler, Wu & Nguyen by having a head broader than long (vs. head longer than wide in L. quangninhensis), tympanum invisible (vs. visible in L. quangninhensis), tibia shorter than thigh (vs. tibia longer than thigh in L. quangninhensis), toes fully webbed (vs. webbing formula I0–0II0–1/3III0–1/3IV1/2–0V in L. quangninhensis), and dorsal surface of tibia and foot with distinct tubercles (vs. small tubercles in L. quangninhensis); from L. taylori Matsui, Panha, Khonsue & Kuraishi by having a head broader than long (vs. head longer than wide in L. taylori), tympanum invisible (vs. slightly visible in L. taylori), length of fingers IL. taylori), and the presence of dorsal ridges (vs. absence in L. taylori); from L. savan Phimmachak, Richards, Sivongxay, Seateun, Chuaynkern, Makchai, Som & Stuart by the absence of caruncle on head in males (vs. presence in L. savan), tympanum invisible (vs. visible in L. savan), relative length of toes IL. savan), and males with nuptial pad on Finger I (vs. absence in L. savan).

Limnonectes phuyenensis sp. nov. (Figs. 2���5) Holotype: IEBR 4737 (Field number DTD 138), adult male, collected by D. T. Do on 28 March 2015 (12��51.184 ��� N, 109��13.151 ��� E, at an elevation of 359 m asl.) in the evergreen forest of Ca Mountain Pass, near Hoa Thinh Commune, Tay Hoa District, Phu Yen Province, Vietnam. Paratypes: 18 specimens collected from the evergreen forest of Phu Yen Province, Vietnam: One adult male, PYU 535 (Field number DTD 04) and two adult females, IEBR 4739, 4740 (Field numbers DTD 163, 171), collected by D. T. Do on 28 March 2015 (12��51.184 ��� N, 109��13.151 ��� E, at an elevation of 359 m asl.) from Ca Mountain Pass, near Hoa Thinh Commune, Tay Hoa District; one adult male, IEBR 4738 (Field number DTD 535) and one adult female, PYU 536 (Field number DTD 536), collected by D. T. Do on 19 August 2015 (12��52.470 ��� N, 108��49.511 ��� E, at an elevation of 554 m asl.) from Ca Mountain Pass, near Son Thanh Tay Commune, Tay Hoa District; one adult male, ZFMK 103169 (Field number DTD 680) and one adult female, ZFMK 103170 (Field number DTD 708), collected by D. T. Do on 10 September 2016 (13��27.437 ��� N, 108��52.005 ��� E, at an elevation of 430 m asl.) from Phu Mo Commune, Dong Xuan District; one adult male, ITBCZ 4598 (Field number SH 040) collected by L. T. Nguyen, M. V. Le, and H. V. Lo on 26 March 2019 (12��78.549 ��� N, 109��03.737 ��� E, at an elevation of 350 m asl.); six adult males, ITBCZ 4746, 4747, 4749, 4753, 4754 (Field numbers SH 188, 189, 191, 195, 196, respectively) and VNMN 010902 (Field number SH194) and four adult females, ITBCZ 4745, 4748, 4750 (Field numbers SH 187, 190, 192, 193, respectively) and VNMN 010903 (Field number SH193) collected by L. T. Nguyen, M. V. Le, and H. V. Lo on 31 March 2019 (12��78.549 ��� N, 109��03.737 ��� E, at an elevation of 350 m asl.) from Song Hinh Protection Forest, Song Hinh District. Diagnosis: Molecular and morphological data, including the presence of fang-like odontoid processes on the lower jaw and sexual dimorphism, with males larger than females and enlargement of the head in males (Emerson et al. 2000), revealed the new species to be nested within Limnonectes. Limnonectes phuyenensis sp. nov. is distinguishable from its congeners by a combination of the following morphological characters: (1) SVL 49.8���77.9 mm in males, 40.8���61.8 mm in females; (2) males with moderately enlarged head (HL/SVL 0.47, HW/SVL 0.48); (3) head slightly broader than long (HL/HW 0.97 in males, 0.95 in females); (4) vomerine teeth present; (5) external vocal sacs absent; (6) rostral length short (RL/SVL 0.15); (7) tympanum invisible; (8) dorsal surface of head, body and flanks with ridges and tubercles; (9) dorsal surface of tibia and foot distinctly tuberculate, covered with moderately dense, small, low tubercles; (10) supratympanic fold present; (11) dorsolateral fold absent; (12) Finger I with nuptial pad, composed of minute spines on dorsal surface and medial edge in males; (13) toes webbed to distal of terminal phalanx, webbing formala I0���0II0���0III0���0IV0 - ���0V0; (14) in life, dorsum yellowish brown with dark brown markings; (15) ventral surface white with brown markings. Description of holotype: Adult male (IEBR 4737); SVL 62.8 mm; habitus robust with enlarged head (HL/SVL 0.48, HW/SVL 0.50); head slightly broader than long (HL 30.1 mm, HW 31.2 mm); snout round anteriorly in dorsal view, projecting beyond lower jaw; nostril lateral, closer to tip of snout than to eye (NS 4.2 mm, EN 4.8 mm); canthus rostralis indistinct; loreal region oblique and slightly concave; rostral length greater than eye diameter (RL 9.0 mm, ED 7.6 mm); interorbital distance wider than internarial distance and upper eyelid width (IOD 6.4 mm, IND 5.3 mm, UEW 3.9 mm); tympanum invisible; vomerine teeth in two oblique ridges; tongue cordiform, notched posteriorly; lower jaw with two tooth-like, odontoid processes robust with very sharp tips (OH 4.0 mm); external vocal sac absent. Forelimbs: Arms short; upper arm length (UAL) 10.8 mm, forearm length (FAL) 26.2 mm; relative finger lengths: IHind limbs: Tibia length shorter than thigh length (FeL 28.8 mm, TbL 27.1 mm), approximately 2.2 times longer than wide (TbW 12.4 mm); tips of toes blunt, slightly round; relative length of toes: ISkin texture in life: Dorsal surface of head, body and flanks with ridges and tubercles; small tubercles on upper eyelid; supratympanic fold distinct, extending from eye to angle of jaw; dorsolateral fold absent; small tubercles around cloaca; dorsal surface of forelimbs and thighs smooth; dorsal surface of tibia and foot distinctly tuberculate, covered with moderately dense, small, low tubercles; throat, chest, belly and ventral surface of thighs smooth. Coloration in life: Iris dark gray in upper half and light gray in lower half; head with a pale yellow bar in anterior interorbital region and a narrow dark brown bar in posterior interorbital region; dorsum and flanks with yellowish brown marking; lips with dark bars; supratympanic fold dark brown; dorsal surface of limbs yellowish brown with dark crossbars; ventral surface of limbs, throat, chest, and belly white with brown markings; toe webbing brown. Coloration in preservative: Dorsum and upper part of flanks pale brown with dark brown marking; lower part of flanks white with brown marking; lips with dark bars; head with a narrow dark brown bar in anterior interorbital region; supratympanic fold dark brown; dorsal surface of limbs pale brown with dark crossbars; ventral surface of limbs, throat, chest, and belly white with brown markings; minute spines on Finger I greyish brown; toe webbing brown. Variation and sexual dimorphism: Measurements and morphological characters of the type series are provided in Table 1 and photographs of paratypes in life are presented in Fig. 6. Males are larger than females (SVL 60.1 �� 8.3 mm, n = 11 males vs. 50.3�� 7.3 mm, n = 8 females). Males have a larger head than females (HL/ SVL 0.47 vs. 0.44, HW / SVL 0.48 vs. 0.46). The male specimens have a nuptial pad on Finger I. The males have robust fanglike odontoid processes, which are less pronounced in the females. The females contained yellowish cream eggs of varying sizes. One juvenile (unvouchered) had a yellowish dorsum with distinct brown marking (Figure 6F). ......continued on the next page Etymology. The specific epithet ��� phuyenensis ��� refers to the type locality of the new species, Phu Yen Province. As common names we suggest Phu Yen Wart Frog (English), ���ch nh���o ph�� y��n (Vietnamese). Ecological notes. Specimens were found between 19:00 and 23:00 in the head-water of small, rocky streams (Fig. 7A). They were found in the water or on the ground of stream banks at elevations above 300 m. Males were sometimes found in small water pools with two or three females nearby. The surrounding habitat was secondary forest of large, medium and small hardwoods mixed with shrubs and vines. Air temperature at the sites was 20.5���26.3 oC and relative humidity was 73���96%. The females collected in March, August, and September contained yellowish cream eggs of varying sizes. Male advertisement calls and tadpoles of the species have not been recorded during our field surveys. Other amphibian species found at the sites included Leptobrachella macrops (Duong, Do, Ngo, Nguyen & Poyarkov), Megophrys elfina (Poyarkov, Duong, Orlov, Gogoleva, Vassilieva, Nguyen, Nguyen, Nguyen, Che & Mahony), Limnonectes poilani (Bourret), Odorrana morafkai (Bain, Lathrop, Murphy, Orlov & Ho), Kurixalus cf. banaensis (Bourret), and Rhacophorus annamensis (Smith). Distribution. Limnonectes phuyenensis sp. nov. is currently known only from Phu Yen Province, South-central Vietnam (Fig. 8). Comparisons. We compared the new species with its congeners from Vietnam and neighboring countries including Laos, Cambodia, Thailand, and China. Limnonectes phuyenensis sp. nov. differs from L. bannaensis Ye, Fei & Jiang by having a smaller size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 74.9 mm, n= 10 males, 60.8 mm, n= 8 females in L. bannaensis), head broader than long (vs. head longer than wide in L. bannaensis), tips of toes blunt (vs. tips of toes expanded into discs in L. bannaensis), dorsal surface of tibia and foot with tubercles (vs. smooth in L. bannaensis), and males with nuptial pad on Finger I (vs. males with nuptial pad on fingers I and II in L. bannaensis); from L. coffeatus Phimmachak, Sivongxay, Seateum, Yodthong, Rujirawan, Neang, Aowphol & Stuart by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 37.9 mm, n= 1 male, 40.5 mm, n= 3 females in L. coffeatus), males with nuptial pad on Finger I (vs. absence in males of L. coffeatus), tympanum invisible (vs. visible in L. coffeatus), and the absence of W marking on head (vs. presence in L. coffeatus); from L. dabanus (Smith) by the absence of caruncle on head in males (vs. presence in L. dabanus), tympanum invisible (vs. visible in L. dabanus), toes fully webbed (vs. webbing formula I0���1/3II0���1/4III0���1IV1���0V in L. dabanus), and males with nuptial pad on Finger I (vs. absence in L. dabanus); from L. fastigatus by having a differrent odontoid processes shape (robust with very sharp tips vs. thin and elongated in L. fastigatus), nostril closer to tip of snout than to eye (vs. midway between snout tip and eye in L. fastigatus), and belly with brown flecking or vermiform markings in preservative (vs. creamy white in preservative in L. fastigatus); from L. fragilis (Liu & Hu) by having a head broader than long (vs. head longer than wide in L. fragilis), a greater ratio of HL/SVL and HW/SVL (HL/SVL=0.47, HW/SVL=0.48, n= 11 males and HL/SVL=0.44, HW/SVL=0.46, n= 8 females vs. HL/SVL=0.45, HW/SVL=0.42, n= 6 males and HL/SVL=0.42, HW/SVL=0.41, n= 10 females in L. fragilis), tibia length shorter than thigh length (vs. tibia length longer than thigh length in L. fragilis), and males with nuptial pad on Finger I (vs. absence in males of L. fragilis); from L. fujianensis Ye & Fei by having a head broader than long (vs. head longer than wide in L. fujianensis), a greater ratio of HL/SVL and HW/SVL (HL/SVL=0.47, HW/SVL=0.48, n= 11 males and HL/SVL=0.44, HW/SVL=0.46, n= 8 females vs. HL/SVL=0.45, HW/SVL=0.44, n= 20 males and HL/SVL=0.38, HW/SVL=0.37, n= 20 females in L fujianensis), males with nuptial pad on Finger I (vs. nuptial pad on Fingers I and II in L. fujianensis), and the absence of W marking on head (vs. presence in L. fujianensis); from L. gyldenstolpei (Andersson) by the absence of caruncle on head in males (vs. presence in L. gyldenstolpei), tympanum invisible (vs. visible in L. gyldenstolpei), toes fully webbed (vs. webbing formula I0���3/4II0���1III1/4���1IV11/2���0V in L. gyldenstolpei), and males with nuptial pad on Finger I (vs. absence in L. gyldenstolpei); from L. kiziriani by having larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 54.1 mm, n= 13 males, 46.8 mm, n= 7 females in L. kiziriani), head broader than long (vs. longer than wide in L. kiziriani), a smaller ratio of TbL/TbW (TbL/TbW 2.31, n= 11 males and 2.40, n= 8 females vs. TbL/TbW 2.54, n= 13 males and 2.57, n= 7 females in L. kiziriani), and the absence of two dermal ridges parallel from posterior of eye to the end of shoulder (vs. presence in L. kiziriani); from L. hascheanus (Stoliczka) by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm n= 8 females vs. 22.1 mm, n= 9 males, 23.0 mm, n= 9 females of L. hascheanus), the absence of dorsolateral folds (vs. presence in L. hascheanus), males with nuptial pad on Finger I (vs. absence in males of L. hascheanus), and tympanum invisible (vs. visible in L. hascheanus); from L. isanensis McLeod, Kelly & Barley by having a smaller size in males (SVL 60.1 mm, n=11 vs. 71.5 mm, n= 11 in L. isanensis), head broader than long (vs. head longer than wide in L. isanensis), dorsum and flank with tubercles (vs. absence in L. isanensis), and males with nuptial pad on Finger I (vs. nuptial pad on Fingers I and II in L. isanensis); from L. jarujini Matsui, Panha, Khonsue & Kuraishi by having a smaller size in males (SVL 60.1 mm, n=11 vs. 73.3 mm, n= 8 in L. jarujini), head broader than long (vs. head longer than wide in L. jarujini), length of fingers IL. jarujini), and dorsum and flank with tubercles (vs. absence in L. jarujini); from L. khammonensis (Smith) by having a larger size in females (SVL 50.3 mm, n=8 vs. 37 mm, n= 1 in L. khammonensis) and tympanum invisible (vs. visible in L. khammonensis); from L. kohchangae (Smith) by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 35.9 mm, n= 21 males, 35.2 mm, n= 14 females of L. kohchangae), tympanum invisible (vs. visible in L. kohchangae) and males with nuptial pad on Finger I (vs. absent in L. kohchangae); from L. limborgi (Sclater) by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 33.0 mm, n= 30 males, 31.7 mm, n= 24 females of L. limborgi), the absence of dorsolateral folds (vs. presence in L. limborgi), males with nuptial pad on Finger I (vs. absence in males of L. limborgi), and tympanum invisible (vs. visible in L. limborgi); from L. liui (Yang) by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 35.7 mm, n= 3 males, 32.7 mm, n= 1 female of L. liui), tympanum invisible (vs. visible in L. liui), and dorsolateral fold absent (vs. present in L. liui); from L. longchuanensis Suwannapoom, Yuan, Sullivan & McLeod by having head broader than long (vs. head longer than wide in L. longchuanensis), males with nuptial pad on Finger I (vs. nuptial pad on Fingers I and II in L. longchuanensis), and dorsal ridges present (vs. absent in L. longchuanensis); from L. lauhachindai Aowphol, Rujirawan, Taksinum, Chuaynkern & Stuart by having a larger size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 33.8 mm, n= 20 males, 35.2 mm, n= 4 females in L. lauhachindai), the absence of caruncle on head (vs. presence in L. lauhachindai), tympanum invisible (vs. visible in L. lauhachindai), and males with nuptial pad on Finger I (vs. absence in L. lauhachindai); from L. macrognathus (Boulenger) by having a larger size in males (SVL 60.1 mm, n=11 vs. 45.5 mm, n= 3 in L. macrognathus), the absence of caruncle on head (vs. presence in L. macrognathus), and tympanum invisible (vs. visible in L. macrognathus); from L. megastomias McLeod by having a smaller size (SVL 60.1 mm, n= 11 males, 50.3 mm in, n= 8 females vs. 80.2 mm, n= 22 males, 74.0 mm, n= 11 females), head broader than long (vs. head longer than wide in L. megastomias), dorsal surface of tibia and foot with tubercles (vs. smooth in L. megastomias), and males with nuptial pad on Finger I (vs. nuptial pad on Fingers I and II in L. megastomias); from L. nguyenorum by having a smaller ratio of HL/SVL (0.47, n= 11 males, 0.44, n= 8 females vs. 0.42���0.43, n= 2 males, 0.39���0.39, n= 2 females in L. nguyenorum), the head broader than long (vs. head longer than wide in L. nguyenorum), tibia shorter than thigh (vs. tibia longer than thigh in L. nguyenorum), males with nuptial pad on Finger I (vs. nuptial pad on Fingers I and II in L. nguyenorum), and the absence of two dermal ridges parallel from posterior of eye to the end of shoulder (vs. presence in L. nguyenorum); from L. poilani (Bourret) by having a smaller size (SVL 60.1 mm, n= 11 males, 50.3 mm, n= 8 females vs. 90.0 mm, n= 4 males, 80.2 mm, n= 6 females of L. poilani), head broader than long (vs. head longer than wide in L. poilani), tibia shorter than thigh (vs. tibia longer than thigh in L. poilani), tympanum invisible (vs. visible in L. poilani), and ventral surface of limbs and abdomen white with brown markings (vs. yellow in L. poilani); from L. quangninhensis Pham, Le, Nguyen, Ziegler, Wu & Nguyen by having a head broader than long (vs. head longer than wide in L. quangninhensis), tympanum invisible (vs. visible in L. quangninhensis), tibia shorter than thigh (vs. tibia longer than thigh in L. quangninhensis), toes fully webbed (vs. webbing formula I0���0II0���1/3III0���1/3IV1/2���0V in L. quangninhensis), and dorsal surface of tibia and foot with distinct tubercles (vs. small tubercles in L. quangninhensis); from L. taylori Matsui, Panha, Khonsue & Kuraishi by having a head broader than long (vs. head longer than wide in L. taylori), tympanum invisible (vs. slightly visible in L. taylori), length of fingers IL. taylori), and the presence of dorsal ridges (vs. absence in L. taylori); from L. savan Phimmachak, Richards, Sivongxay, Seateun, Chuaynkern, Makchai, Som & Stuart by the absence of caruncle on head in males (vs. presence in L. savan), tympanum invisible (vs. visible in L. savan), relative length of toes IL. savan), and males with nuptial pad on Finger I (vs. absence in L. savan)., Published as part of Pham, Cuong The, Do, Dang Trong, Le, Minh Duc, Ngo, Hanh Thi, Nguyen, Luan Thanh, Ziegler, Thomas & Nguyen, Truong Quang, 2020, A new species of Limnonectes (Amphibia: Anura: Dicroglossidae) from Vietnam, pp. 387-402 in Zootaxa 4894 (3) on pages 390-398, DOI: 10.11646/zootaxa.4894.3.5, http://zenodo.org/record/4315907, {"references":["Emerson, S., Inger, R. & Iskandar, D. (2000) Molecular systematics and biogeography of the fanged frogs of Southeast Asia. Molecular Phylogenetics and Evolution, 16, 131 - 142. https: // doi. org / 10.1006 / mpev. 2000.0778"]}

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 &dstrok;i���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]"]}

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), Rắn xe &dstrok;iếu tràng 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 (Günther).

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 houaphanensis sp. nov. (Fig. 2) Holotype. One adult male, IEBR A.2013.109, collected by Alexandre Teyni�� 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, R��sler, 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., 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 507-516, DOI: 10.11646/zootaxa.4822.4.3, http://zenodo.org/record/4452362, {"references":["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.","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","David, P., Teynie, A. & Ohler, A. (2004) A new species of Cyrtodactylus Gray, 1827 (Reptilia: Squamata: Gekkonidae) from southern Laos. The Raffles Bulletin of Zoology, 52 (2), 621 - 627.","Teynie, A. & David, P. (2010) Voyages naturalistes au Laos. Les Reptiles. Editions Revoir, Nohanent, 315 pp.","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","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.","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","Stuart, B. L. (1999) Amphibians and reptiles. In: Duckworth, J. W., Salter, R. E. & Khounboline, K. (Eds.), Wildlife in Lao PDR: 1999 status report. IUCN, WCS & CPAWM, Ventiane, pp. 43 - 67.","Luu, V. Q., 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","Ngo, V. T. & Pauwels, O. S. G. (2010) A new cave-dwelling species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Khammouane Province, southern Laos. Zootaxa, 2730 (1), 44 - 56. https: // doi. org / 10.11646 / zootaxa. 2730.1.3","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., Nguyen, T. Q., Schmitz, A., Kingsada, P., Auer, M. & T. Ziegler (2011) A new species of karst dwelling Cyrtodactylus (Squamata: Gekkonidae) from northwestern Laos. Zootaxa, 2930 (1), 1 - 21. https: // doi. org / 10.11646 / zootaxa. 2930.1.1","Rosler, H., Vu, T. N., Nguyen, T. Q., Ngo, T. V. & Ziegler, T. (2008) A new Cyrtodactylus (Squamata: Gekkonidae) from central Vietnam. Hamadryad, 33, 48 - 63.","Luu, V. Q., Calame, T., Nguyen, T. Q., Soudthichak, S., Bonkowski, M. & Ziegler, T. (2013) New country records of reptiles from Laos. Biodiversity Data Journal, 1, 1 - 14. https: // doi. org / 10.3897 / BDJ. 1. e 1015","Luu, V. Q., Calame, T., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016 b) Cyrtodactylus rufford, a new cave-dwelling 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., 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","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.","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 (1), 29 - 40. https: // doi. org / 10.11646 / zootaxa. 2833.1.3","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), 80 - 96. https: // doi. org / 10.11646 / zootaxa. 3835.1.4","Nguyen, T. Q., Kingsada, P., Rosler, H., Auer, M. & Ziegler, T. (2010) A new species of Cyrtodactylus (Squamata: Gekkonidae) from northern Laos. Zootaxa, 2652 (1), 1 - 16. https: // doi. org / 10.11646 / zootaxa. 2652.1.1"]}

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"]}

Cyrtodactylus ngoiensis sp. nov. (Fig. 4) Holotype. One adult male, IEBR 4548 (Fig 4a), collected by Alexandre Teynié 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 Teynié 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).

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

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).

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 Th���ch s��ng d���p 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 25��C to 31��C 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)., Published as part of Do, Quyen Hanh, Pham, Cuong The, Phan, Tien Quang, Le, Minh Duc, Ziegler, Thomas & Nguyen, Truong Quang, 2020, A new species of Hemiphyllodactylus (Squamata: Gekkonidae) from Tuyen Quang Province, Vietnam, pp. 511-532 in Zootaxa 4821 (3) on pages 515-517, DOI: 10.11646/zootaxa.4821.3.5, http://zenodo.org/record/4401075, {"references":["Boulenger, G. A. (1903) Descriptions of new lizards in the collection of the British Museum. The Annals and Magazine of Natural History, Series 7, 12 (70), 429 - 435. https: // doi. org / 10.1080 / 00222930308678877","Barbour, T. (1924) A Yunnan gecko. Occasional Papers of the Boston Society of Natural History, 5, 133 - 135.","Smith, M. A. (1935) The Fauna of British India, including Ceylon and Burma. Reptiles and Amphibia. Vol. II. Sauria. Taylor and Francis, London, 440 pp.","Taylor, E. H. (1963) The lizards of Thailand. The University of Kansas Science Bulletin, 44, 687 - 1077.","Zhou, K. - Y., Liu, Y. - Z. & Yang, G. - P. (1981) Three new subspecies of Hemiphyllodactylus yunnanensis (Boulenger) from China. Acta Zootaxonomica Sinica, 6, 202 - 209. [in Chinese, English translation by H. Ota (1996) in Smithsonian Herpetological Information Service, 110, 8 pp., 1 pl.]","Bourret, R. (2009) Les lezards de l'Indochine. Edition Chimaira, Frankfurt am Main, 624 pp.","Zug, G. R. (2010) Speciation and dispersal in a low diversity taxon: The Slender Geckos Hemiphyllodactylus (Reptilia, Gekkonidae). Smithsonian Contributions to Zoology, 631, 1 - 70. https: // doi. org / 10.5479 / si. 00810282.631","Grismer, L. L., Wood, P. L. Jr., Anuar, S., Muin, M. A., Quah, E. S. H. & Mcguire, J. A. (2013) Integrative taxonomy uncovers high levels of cryptic species diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the description of a new species from Peninsular Malaysia. Zoological Journal of the Linnean Society, 169, 849 - 880. https: // doi. org / 10.1111 / zoj. 12064","Grismer, L. L., Riyanto, A., Iskandar, D. T. & McGuire, J. A. (2014 a) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Pulau Enggano, southwestern Sumatra, Indonesia. Zootaxa, 3821 (4), 485 - 495. https: // doi. org / 10.11646 / zootaxa. 3821.4.7","Nguyen, T. Q., Lehmann, T., Le, M. D., Duong, H. T., Bonkowski, M. & Ziegler, T. (2013) A new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Vietnam. Zootaxa, 3736 (1), 89 - 98. https: // doi. org / 10.11646 / zootaxa. 3736.1.5","Nguyen, T. Q., Botov, A., Le, M. D., Nophaseud, L., Zug, G., Bonkowski, M. & Ziegler, T. (2014) A new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Laos. Zootaxa, 3827 (1), 45 - 56. https: // doi. org / 10.11646 / zootaxa. 3827.1.4","Nguyen, T. Q., Do, Q. H., Ngo, H. T. Pham, A. V., Pham, C. T., Le, M. D. & Ziegler, T. (2020) Two new species of Hemiphyllodactylus (Squamata: Gekkonidae) from Hoa Binh Province, Vietnam. Zootaxa, 4801 (3), 513 - 536. https: // doi. org / 10.11646 / zootaxa. 4801.3.5","Ngo, T. V., Grismer, L. L., Pham, T. H. & Wood, P. L. Jr. (2014) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Ba Na-Nui Chua Nature Reserve, Central Vietnam. Zootaxa, 3760 (4), 539 - 552. https: // doi. org / 10.11646 / zootaxa. 3760.4.3","Guo, W., Zhou, K., Yan, J. & Li, P. (2015) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from western Yunnan, China. Zootaxa, 3974 (3), 377 - 390.","Cobos, A. L., Grismer, L. L., Wood, P. L. Jr, Quah, E. S. H., Anuar, S. & Muin, M. A. (2016) Phylogenetic relationships of geckos of the Hemiphyllodactylus harterti group, a new species from Penang Island, Peninsular Malaysia, and a likely case of true cryptic speciation. Zootaxa, 4107 (3), 367 - 380. https: // doi. org / 10.11646 / zootaxa. 4107.3.5","Yan, J., Lin, Y., Guo, W., Li, P. & Zhou, K. (2016) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Guizhou, China. Zootaxa, 4117 (4), 543 - 554. https: // doi. org / 10.11646 / zootaxa. 4117.4.6","Sukprasert, A., Sutthiwises, S., Lauhachinda, V. & Taksintum, W. (2018) Two new species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from Thailand. Zootaxa, 4369 (3), 363 - 376. https: // doi. org / 10.11646 / zootaxa. 4369.3.4","Eliades, S. J., Phimmachak, S., Sivongxay, N., Siler, C. D. & Stuart, B. L. (2019) Two new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from Laos. Zootaxa, 4577 (1), 131 - 147. https: // doi. org / 10.11646 / zootaxa. 4577.1.8","Agarwal, I., Khandekar, A., Giri, V. B. & Ramakrishnan, U. (2019) The hills are alive with geckos! A radiation of a dozen species on sky islands across peninsular India (Squamata: Gekkonidae, Hemiphyllodactylus) with the description of three new species, Organisms Diversity & Evolution, 19 (2), 341 - 361. https: // doi. org / 10.1007 / s 13127 - 019 - 00392 - 5","Agarwal, I., Bauer, A. M., Pal, S. N., Srikanthan, A. N. & Khandekar, A. (2020) Two more new Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Tamil Nadu, India. Zootaxa, 4729 (2), 249 - 265. https: // doi. org / 10.11646 / zootaxa. 4729.2.6"]}

Zhangixalus franki sp. nov. urn:lsid:zoobank.org:act: 346FA039-3D9A-4F41-A4F2-57F5E7ABA793 Fig. 2 Diagnosis The new species can be assigned to the genus Zhangixalus based on the following morphological characters: snout pointed; presence of intercalary cartilage between terminal and penultimate phalanges of digits, distal end of terminal phalanx in Y-shape; tips of digits expanded into large disks, bearing circum-marginal grooves; fingers webbed; presence of supra-cloacal dermal ridge; and pupil horizontal (Jiang et al. 2019). In addition, the new species is unambiguously nested in the genus Zhangixalus by molecular phylogenetic evidence. It is closely related to Z. duboisi and Z. omeimontis and nested in the same clade with Zhangixalus dugritei (David, 1872), the type species of this genus. The new species is distinguished from its congeners by a combination of the following morphological characters: 1) size large (SVL 77.9���85.8 mm for the males); 2) head slightly wider than long; 3) snout pointed; 4) dorsal skin smooth; 5) finger webbing formula I1-1II0-1III��-0IV; 6) dorsal surface of head and body green with dark brown spots; 7) lower jaw region greyish, throat, chest and belly white; 8) white stripe along edge of jaw, insertion of limbs, and along lateral ridges of fore limbs, hind limbs and flank, separating upper green part from lower grey part. Etymology We name this new species in honor of Dr Frank Mutschmann, late amphibian veterinarian and conservationist from Berlin, Germany, in recognition of his support of our amphibian research and conservation projects in Vietnam. As common names we suggest Frank���s Tree Frog (English), ���ch c��y frank (Vietnamese) and Franks Ruderfrosch (German). Material examined Holotype VIETNAM ��� adult &male;; northern Vietnam, Ha Giang Province, Quan Ba District, in the forest of Tung Vai Commune; 23002���45��� N, 104050���59��� E; elevation 1360 m a.s.l.; 11 Jun. 2019; collected by T.T. Nguyen; VNMN 011682. Paratypes VIETNAM ��� 4 adult &male;&male;; same data as for holotype; 13 Jun. 2019; collected by T.T. Nguyen; VNMN 011683, VNMN 011684, IEBR A.2019.7-A.2019.8 (VNMN 011685, VNMN 011686) and one sub-adult VNMN 011687. Description of holotype SIZE. Large, body robust (SVL 82.8 mm). HEAD. Slightly compressed, wider than long (HW 26.5 mm, HL 25.7 mm), convex above; snout pointed, slightly protruding beyond lower jaw in lateral view, and longer than horizontal diameter of eye (SNL 11.6 mm, ED 8.2 mm). CANTHUS ROSTRALIS. Rounded, loreal region oblique, concave; interorbital distance greater than internarial distance and upper eyelid width (IOD 9.2 mm, IN 8.1 mm, UEW 6.2 mm); distance between anterior corners of eyes approximately 69% of distance between posterior corners of eyes. NOSTRILS. Round, without lateral flap of skin, closer to tip of snout than to eye. PUPIL. Oval, horizontal. TYMPANUM. Distinct, round, about half of eye diameter, and two times greater than distance between tympanum and eye. PINEAL OCELLUS. Absent; spinules on upper eyelid absent; vomerine teeth well developed, in two oblique ridges. CHOANAE. Round. TONGUE. Deeply notched posteriorly; supratympanic fold distinct, extending from behind eye to beyond level of axilla. FORELIMBS. Upper arm short, about one third of forearm length (FLL 14.7 mm, HAL 41.7 mm), dermal fringe present along outer edge of forearm; not well developed; relative finger lengths IZhangixalus distributed in Vietnam and neighboring countries. Zhangixalus franki sp. nov can be distinguished from the following 29 species in the genus Zhangixalus by having a larger size in males (SVL 77.9-85.8) versus Z. achantharrhena at 36.52���40.56 mm (Harvey et al. 2002); Z. arboreus at 42���60 mm (Wilkinson 2003); Z. arvalis at 39���46.4 mm (Lue et al. 1995); Z. aurantiventris at 48���54 mm (Lue et al. 1994); Z. chenfui at 33���41 mm (Fei et al. 2010); Z. dorsoviridis (Bourret, 1937) at 31.31��� 2.43 mm (Orlov et al. 2001); Z. dugritei at 31.65���47 mm (Liu 1950; Orlov et al. 2001; Fei et al. 2010); Z. dulitensis at 38.39���39.66 mm (Harvey et al. 2002); Z. hongchibaensis at 46.5���49.7 mm (Li et al. 2012a); Z. hungfuensis at 31���37 mm (Fei et al. 2010) Z. jarujini at 33.7��� 40 mm (Fei et al. 2010); Z. leucofasciatus at 35���48 mm (Fei et al. 2010); Z. lishuiensis at 34.2���35.8 mm (Liu et al. 2017); Z. minimus at 21.3���33.0 mm (Rao et al. 2006); Z. moltrechti at 33���46 mm (Fei et al. 2010); Z. nigropunctatus at 32���37 mm (Fei et al. 2010); Z. owstoni at 42���51 mm (Maeda & Matsui 1990); Z. pinglongensis at 32.0��� 38.5 mm (Mo et al. 2016); Z. prasinatus at 49���56 mm (Fei et al. 2010); Z. prominanus at 50.46���51.26 mm (Harvey et al. 2002); Z. puerensis at 35.5���41 mm (Bain & Nguyen 2004); Z. schlegelii at 49 mm (Stejneger 1907); Z. suffry at 38.5���52.9 mm (Bordoloi et al. 2007); Z. taipeianus at 30.7���36.7 mm (Liang & Wang 1978); Z. viridis at 45���56 mm (Goris & Maeda 2004); Z. wui at 35.2���38.2 mm (Li et al. 2012a); Z. yaoshanensis at 31.6���36.4 mm (Chen et al. 2018); Z. yinggelingensis at 43���43.4 mm (Chou et al. 2007); Z. zhoukaiyae at 27.9���36.88 mm (Pan et al. 2017). Zhangixalus franki sp. nov. differs from the following seven species by having dorsum smooth; dorsal surface green with some small dark spots; chest and belly grey; supratympanic fold well developed, brown or gold; white stripe along lateral ridges of fore and hind limbs and flank, separating upper green part from lower grey part; and webbing between fingers incomplete verses dorsum of Z. duboisi green and brown, skin granular with horny spinules, venter fleshy with brown spots (Ohler et al. 2000); snout of Z. hui yellowish brown (Li et al. 2012a); dorsum and dorsal aspect of limbs of Z. omeimontis green with large brown markings (Liu 1950), supratympanic fold of Z. pachyproctus, Z. smaragdinus and Z. dennysi green and weakly developed (Yu et al. 2019; Fei et al. 2010); lateral ridges of fore and hind limbs and flank of Z. dennysi without white stripe; supratympanic fold yellow and webbing between fingers complete in Z. feae (Ziegler et al. 2014). The new species mostly resembles Z. burmanus, but with clearly distinguishing characters: males of Zhangixalus franki sp. nov. with SVL at 77.9���85.5 mm larger than those of Z. burmanus with SVL at 47���70.5 (Ohler 2009; Jiang et al. 2016; Wilkinson & Rao 2004); webbing between fingers in males of Zhangixalus franki sp. nov. is more developed than that in males of Z. burmanus (I1-1II0-1III��-0IV vs I1���1��II1���2��III1���1IV in Z. burmanus Ohler 2009); color on upper and, lower parts of flank separated from each other by white line, and dark spots on anterior and posterior sides of thigh are absent in Zhangixalus franki sp. nov. whereas, dark brown spots enclosing cream spots on flanks and thigh present in Z. burmanus (Wilkinson & Rao 2004). Distribution Zhangixalus franki sp. nov. is currently known only from the type locality in Ha Giang Province, northern Vietnam (Fig. 3). The species was recorded at elevations between 1320 and 1360 m a.s.l. Natural history Specimens of the new species were collected from 19:00 to 24:00 on trees, close to small ponds in undisturbed evergreen forest (Fig. 4). Other tree frogs that were found at the same site were Polypedates sp. (of the P. leucomystax species complex), Z. duboisi and Kurixalus sp. Females, larval stages and eggs of the new species are unknown. Conservation status The new species is expected to be found in evergreen forests of Guangxi Province, southern China. However, the actual distributional range should be confirmed in further studies. Given the available information, we suggest this species be considered as Data Deficient following IUCN���s Red List categories (IUCN 2020)., Published as part of Ninh, Hoa Thi, Nguyen, Tao Thien, Orlov, Nikolai, Nguyen, Truong Quang & Ziegler, Thomas, 2020, A New Species of the Genus Zhangixalus (Amphibia: Rhacophoridae) from Vietnam, pp. 1-18 in European Journal of Taxonomy 688 on pages 7-12, DOI: 10.5852/ejt.2020.688, http://zenodo.org/record/3961720, {"references":["Jiang D., Jiang K., Ren J., Wu J. & Li J. 2019. Resurrection of the genus Leptomantis, with description of a new genus to the family Rhacophoridae (Amphibia: Anura). Asian Herpetological Research 10: 1 - 12. https: // doi. org / 10.16373 / j. cnki. ahr. 180058","Harvey M. B., Pemberton A. J. & Smith E. N. 2002. New and poorly known parachuting frogs (Rhacophoridae: Rhacophorus) from Sumatra and Java. Herpetological Monographs 16 (1): 46 - 92. https: // doi. org / 10.1655 / 0733 - 1347 (2002) 016 [0046: NAPKPF] 2.0. 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Zhangixalus dugritei CHINA • 2 &male;; Sichuan; CIB 20050194, KUHE 27701., Published as part of Ninh, Hoa Thi, Nguyen, Tao Thien, Orlov, Nikolai, Nguyen, Truong Quang & Ziegler, Thomas, 2020, A New Species of the Genus Zhangixalus (Amphibia: Rhacophoridae) from Vietnam, pp. 1-18 in European Journal of Taxonomy 688 on page 18, DOI: 10.5852/ejt.2020.688, http://zenodo.org/record/3961720