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Genus Copris Geoffroy, 1762 Type species: Scarabaeus lunaris Linnaeus, 1758 (subsequent designation by Latreille 1810), Published as part of Bui, Van Bac, Ziegler, Thomas & Bonkowski, Michael, 2019, Checklist of beetles in the subgenus Copris (Paracopris) Balthasar from Asia with description of a new species, and redescription of Copris (Paracopris) punctulatus Wiedemann (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 51-64 in Zootaxa 4712 (1) on page 54, DOI: 10.11646/zootaxa.4712.1.3, http://zenodo.org/record/3586826

Bui, Van Bac, Ziegler, Thomas, Bonkowski, Michael (2019): Checklist of beetles in the subgenus Copris (Paracopris) Balthasar from Asia with description of a new species, and redescription of Copris (Paracopris) punctulatus Wiedemann (Coleoptera: Scarabaeidae: Scarabaeinae). Zootaxa 4712 (1): 51-64, DOI: https://doi.org/10.11646/zootaxa.4712.1.3

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

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Lycodon banksi sp. nov. Figs 2-5, Table 3 Holotype: VNUF R.2015.20 (field number: TK 20.15), adult male, collected on 4 April 2015 by Vinh Quang Luu and Thomas Calame in the karst forest, at the mouth of a cave, Phou Hin Poun NPA, Hinboun District, Khammouane Province, central Laos, at an elevation of 167 m a.s.l. Diagnosis: Lycodon banksi sp. nov is characterized by the following morphological characters: (1) dorsal scales in 17-17-15 rows, dorsal scales on the anterior 2/3 of the body length smooth, the six central dorsal scale rows of the posterior 1/3 of the body length feebly keeled; (2) supralabials 8; (3) infralabials 10; (4) loreal entering orbit; (5) cloacal single; (6) ventral scales 241; (7) dorsal surface of body with 87 greyish yellow blotches; (8) ventral surface of body and tail uniformly grey cream. Description of the holotype: Head elongate (HL 15.3 mm), moderately distinct from the neck, longer than wide (HW/HL ratio 0.71), depressed (HH/HL ratio 0.40), narrow anteriorly (IN/IO ratio 0.65); snout elongate (SnL/HL ratio 0.39); nostril lateral, oval shaped, located in the middle of the nasal; eye large (ED/HL ratio 0.17), pupils vertically elliptic; rostral triangular, much broader than high, hardly visible from above; nasal divided into two scales by a vertical ridge along posterior edge of nostril; two square internasals, as wide as long, bordered by two large, subpentagonal prefrontals posteriorly; frontal single, enlarged, pentagonal, narrowed posteriorly; parietals longer than wide, in contact with each other medially, with upper anterior and posterior temporals, paraparietal laterally and four nuchal scales posteriorly; loreal 1/1, elongate, entering orbit; supralabials 8/8, first and second in contact with nasal, third to fifth entering orbit, sixth largest; infralabials 10/10, first pair in broad contact with each other, first to fifth in contact with first pair of chin shields; first and second pairs of chin shields elongate, of the same size and shape, separated by a medial groove, first pair larger than the second; preocular 1/1; postoculars 2/2, of the same size, bordering anterior temporals; anterior temporals 2/2, posterior temporals 3/3, upper ones smaller than lower ones. Body elongate, SVL 415 mm; TaL> 50 mm (tail tip lost); preventrals 2, ventrals 241; subcaudals 26 (tail tip lost), divided, weakly notched laterally; cloacal single; DSR 17-17-15; dorsal scales on the anterior 2/3 of the body length smooth, the six central dorsal scale rows of the posterior 1/3 of the body length feebly keeled; the vertebral scales not enlarged. Colouration in life: Head dark grey, without vertical light nuchal band; dorsal surface of body dark greyyellow with 87 greyish yellow irregular dorsal blotches; first body blotch starting at ventral scale 13, a half vertebral scale covered by this blotch; two yellow stripes on each side, from behind the neck to vent, indistinct posteriorly; ventral scales grey cream; dorsal surface of tail with at least eleven greyish yellow tail blotches, ventral surface of tail grey cream. Hemipenis: The left hemipenis is only in part everted but shows a spinose ornamentation. Additional specimen: One specimen which was not collected but detected and photographed on 22 July 2016 by an arachnology team consisting of Peter Jaeger, Aloke Sahu and Jonas Ewert, in Khammouane Province, in ca. 12.4 km distance from the type locality. The color pattern of this specimen resembles closely that of the holotype. Comparisons: In our phylogenetic analysis, Lycodon banksi sp. nov. is nested in a clade containing L. rufozonatus, L. semicarinatus (Cope), ‘ L. flavozonatus’, L. futsingensis and L. meridionalis. The new species differs from the similar L. meridionalis by having loreal entering the orbit (versus separated from the orbit), dorsal scales on the anterior 2/3 of the body length smooth, the six central dorsal scale rows on the posterior body third feebly keeled (versus distinctly keeled), dorsal head pattern uniform dark grey (versus with yellow-black marbling in L. meridionalis), and ventral surface grey cream (versus yellow with dark spots posteriorly) (see Bourret, 1935; Orlov & Ryabov, 2004); from L. rufozonatus by having loreal entering the orbit (versus usually separated), a distinctly higher ventral scale count (241 versus 185-204), dorsal scales feebly keeled in the posterior body part (versus all smooth), dorsal head pattern uniform dark grey (versus dark brown with yellow borders), and body pattern blotched (versus banded) (Boulenger 1893); from L. semicarinatus by having loreal touching the orbit (versus separated), a higher ventral scale count (241 versus 211-234), dorsal scale rows keeled along posterior 1/3 (versus keeled along anterior half), belly pattern uniform grey cream (versus yellow), and body pattern blotched (versus banded) (Boulenger 1893); from L. flavozonatus by having loreal in contact with the orbit (versus separated), cloacal single (versus divided), six dorsal scale rows on the posterior third of the body feebly keeled (versus 10-12 keeled dorsal scale rows at midbody), dorsal head dark grey (versus black with light markings), and belly pattern uniform grey cream (versus yellow with large black spots); from L. futsingensis by having loreal entering the orbit (versus separated), a higher ventral scale count (241 versus 193-203 in males), dorsal scales feebly keeled in the posterior body part (versus all smooth), and body pattern blotched (versus banded) (Vogel et al., 2012; Neang et al., 2014) (Table 4). The new species has a loreal entering the orbit and thus differs from the following species and subspecies of the Lycodon ruhstrati group which have the loreal separated from the orbit: L. cardamomensis Daltry & Wüster, 2002, L. davidi, L. multifasciatus (Maki, 1931), L. ophiophagus Vogel, David, Pauwels, Sumontha, Norval, Hendrix, Vu & Ziegler, 2009, L. paucifasciatus Rendahl in Smith, 1943, L. ruhstrati ruhstrati (Fischer, 1886), and Lycodon ruhstrati abditus (Vogel et al., 2009). In addition, the new species differs from L. cardamomensis by having more ventral scales (241 versus 215), and in body pattern (87 blotches versus 12 bands); from L. davidi by having more ventral scales (241 versus 224), six dorsal scale rows on the posterior third of the body feebly keeled (versus dorsal scale rows at midbody slightly keeled, outermost rows entirely smooth throughout body), and belly pattern uniform grey cream (versus anterior third whitishcream, posterior part heavily speckled with dark dots); from L. multifasciatus by having more ventral scales (241 versus maximum 237), and dorsal pattern blotched (versus banded); from L. ophiophagus by having more ventral scales (241 versus 211), and dorsal pattern (87 blotches versus 21-22 bands); from L. paucifasciatus by having fewer dorsal scale rows at neck (17 versus 19), more ventral scale rows (241 versus 221-222), six dorsal scale rows on the posterior third of the body feebly keeled (versus two upper rows plus vertebral row distinctly keeled), and dorsal pattern blotched (versus banded) (Neang et al., 2014); from L. r. ruhstrati and Lycodon ruhstrati abditus by having more ventral scales (241 versus 211-228; 241 versus 206-224, respectively), and dorsal pattern blotched (versus banded in the latter) (Vogel et al., 2012); from L. synaptor by having much more ventral scale rows (241 versus 201-203), dorsal pattern with 87 blotches (versus 30-31 bands), and belly pattern uniform grey cream (versus banded) (Vogel & David 2010); from L. zoosvictoriae by having more ventral scales (241 versus 213), dorsal pattern consisting of 87 blotches (versus 31), and having six dorsal scale rows on the posterior third of the body feebly keeled (versus all weakly keeled) (Neang et al., 2014). From the remaining species occurring in Laos, the new species can be distinguished as follows: from L. capucinus by having more ventrals (241 versus 182-211), fewer supralabials (8/8 versus 9-10), cloacal single (versus divided), dorsal blotches 87 (versus reticulated), and greyish yellow blotched body pattern (versus reticulated); from L. fasciatus by having more ventral scale rows (241 versus 182-225), dorsal pattern consisting of 87 blotches (versus 19-49 bands), and belly pattern uniform grey cream (versus white with dark blotches) (Neang et al., 2014); from L. laoensis by having loreal in contact with the orbit (versus separated), more ventrals (241 versus 169-192), and dorsal scales feebly keeled in the posterior body part (versus all smooth) (Neang et al., 2014); from L. septentrionalis by having more infralabials (10 versus 7-8), more ventral scales (241 versus 202-217), and dorsal pattern blotched (versus banded), as well as belly pattern uniform grey cream (versus white) (Neang et al., 2014); from L. subcinctus by the presence of preocular scale (versus absent), having cloacal scale single (versus divided), dorsal pattern blotched (versus banded in anterior part), and more ventral scale rows (241 versus 129-230) (Neang et al., 2014). From the remaining species in the fasciatus group, the new species differs as follows: from L. butleri Boulenger by having more ventral scale rows (241 versus 220-227), dorsal pattern blotched (versus banded), and belly pattern uniform grey cream (versus banded & spotted) (Grismer et al., 2014); from L. cavernicolus by having dorsal head uniformly dark grey (versus light brown), fewer supralabials (8 versus 9 or 10), more dorsal blotches (87 versus 36-45 bands), dorsal scales on the anterior 2/3 of the body length smooth, the six central dorsal scale rows of the posterior 1/3 of the body length feebly keeled (versus all keeled), and greyish yellow blotched pattern on the body (versus white bands); from L. gongshan by having six dorsal scale rows on the posterior third of the body feebly keeled (versus upper and vertebral dorsal rows keeled), more ventral scale rows (241 versus 210- 216), and dorsal pattern with 87 blotches (versus 32- 40 bands) (Vogel & Luo, 2011); from L. liuchengchaoi by having cloacal scale single (versus divided), dorsal pattern consisting of 87 irregular greyish yellow dorsal blotches (versus 40 well-defined yellow rings), and more ventral scales (241 versus 204) (Zhang et al., 2011). Distribution: Lycodon banksi sp. nov. is currently known only from the type locality in the Phou Hin Poun NPA, Khammouane Province, central Laos (Fig. 6). Etymology: The name of the species is dedicated to our friend and colleague Chris Banks, International Coordinator, Philippine Crocodile National Recovery Team, Zoos Victoria, Australia, for his outstanding contributions towards amphibian and reptile conservation, in particular of the Philippine Crocodile. We propose the following common names: Banks’ Wolf Snake (English), Banks Wolfszahnnatter (German). Natural history: The holotype was found at 20:39 h, crawling on a limestone outcrop in the karst forest, approximately 0.3 m above the forest floor, at an elevation of 167 m a.s.l. The humidity at the time of collection was approximately 85% and the air temperature ranged from 23 to 26 oC (Fig. 7). Another specimen was observed 12.4 km away from the type locality, active on the ground at 23:30 h, near a limestone cliff in the secondary forest., Published as part of Luu, Vinh Quang, Bonkowski, Michael, Nguyen, Truong Quang, Le, Minh Duc, Calame, Thomas & Ziegler, Thomas, 2018, A new species of Lycodon Boie, 1826 (Serpentes: Colubridae) from central Laos, pp. 263-276 in Revue suisse de Zoologie 125 (2) on pages 266-271, DOI: 10.5281/zenodo.1414221, {"references":["Bourret R. 1935. Notes herpetologiques sur l'Indochine francaise. IX. Les Serpents de Chapa. Bulletin general de l'Instruction publique, 14 (1934 - 1935), 7 (Mars), 239 - 251 [Reprint: 5 - 17].","Orlov N. & Ryabov S. A. 2004. Revalidization and change of taxonomic status of Dinodon rufozonatus meridionale Bourret, 1935 (Serpentes: Colubridae: Colubrinae). Russian Journal Herpetology 11 (3): 181 - 197.","Boulenger G. A. 1893. Catalogue of the Snakes in the British Museum (Natural History). Volume I. Trustees of the British Museum, London, xiii + 448 pp., 28 plates.","Vogel G., Nguyen T. Q., Kingsada P. & Ziegler T. 2012. A new species of the genus Lycodon Boie, 1826 from Laos (Squamata: Colubridae). North-Western Journal of Zoology 8: 344 - 352.","Neang T., Hartmann T., Hun S., Souter N. J. & Furey N. M. 2014. A new species of wolf snake (Colubridae: Lycodon Fitzinger, 1826) from Phnom Samkos Wildlife Sanctuary, Cardamom Mountains, southwest Cambodia. Zootaxa 3814 (1): 068 - 080. http: // dx. doi. org / 10.11646 / zootaxa. 3814.1.3","Daltry J. C. & Wuster W. 2002. A new species of wolf snake (Serpentes: Colubridae: Lycodon) from the Cardamom Mountains, southwestern Cambodia. Herpetologica, 58 (4): 498 - 504.","Maki M. 1931. A monograph of the snakes of Japan. Dai-ichi Bhobo, Tokyo: 247 pp.","Vogel G., David P., Pauwels O. S. G., Sumontha M., Norval G., Hendrix R., Vu N. T. & Ziegler T. 2009. A revision of Lycodon ruhstrati (Fischer 1886) auctorum (Squamata Colubridae), with the description of a new species from Thailand and a new subspecies from the Asian mainland. Tropical Zoology 22: 131 - 182.","Smith M. A. 1943. The fauna of British India, Ceylon and Burma, including the whole of the Indo-chinese subregion. Reptilia and Amphibia. Vol. III. Serpentes. Taylor & Francis, London, xii + 583 pp.","Fischer J. G. 1886. Herpetologische Notizen. Abhandlungen des Naturwissenschaftlichen Vereins in Hamburg 9: 51 - 67 (1 - 19).","Vogel G. & David P. 2010. A new species of the genus Lycodon (Boie, 1826) from Yunnan Province, China (Serpentes: Colubridae). Bonn zoological Bulletin 57 (2): 289 - 296.","Grismer L. L., Quah E. S. H., Anuar S., Muin M. A., Wood P. L. Jr. & Nor S. A. M. 2014. A diminutive new species of cavedwelling Wolf Snake (Colubridae: Lycodon Boie, 1826) from Peninsular Malaysia. Zootaxa 3815 (1): 051 - 067. http: // dx. doi. org / 10.11646 / zootaxa. 3815.1.3","Vogel G. & Luo J. 2011. A new species of the genus Lycodon (Boie, 1826) from the southwestern mountains of China (Squamata: Colubridae). Zootaxa 2807: 29 - 40.","Zhang J., Jiang K., Vogel G. & Rao D. 2011. A new species of the genus Lycodon (Squamata, Colubridae) from Sichuan Province, China. Zootaxa 2982: 59 - 68."]}

Synapsis tridens Sharp, 1881 Synapsis tridens Sharp, 1881: xcii (original description). Type locality. India, Assam. Material examined. VIETNAM: CAO BANG PROVINCE: Pia Oac Nature Reserve, primary forest, baited pitfall trap, 30.iv–15.v.2016, 22°34′3.1″N 105°53′3.6″E, 1227 m, 1 ♁, 22°33′59.7″N 105°52′48.5″E, 1165 m, 1 ♀, 22°34′3.1″N, 105°53′4.7″E, 1220 m, 1 ♁, 22°34′3.1″N 105°53′4.4″E, 1213 m, 1♀, Van Bac Bui leg. (all in VNFU). THANH HOA PROVINCE: Pu Luong Nature Reserve, primary forest, baited pitfall trap, 5.–25.iv.2016, 20°28′55.1″N 105°14′29.3″E, 958 m, 1♁, 20°28′54.7″N 105°14′30.9″E, 950 m, 1 ♁ 1 ♀, Bùi Văn Bắc leg. (VNUF). Distribution. SW China, NE India, Laos, Myanmar, Thailand and N Vietnam (ZÍDEK & POKORNÝ 2010). Remarks. The aforementioned specimens represent additional records from Vietnam., Published as part of Bui, Van Bac & Bonkowski, Michael, 2018, Synapsis puluongensis sp. nov. and redescription of S. horaki (Coleoptera: Scarabaeidae), with a key to Vietnamese species, pp. 407-418 in Acta Entomologica Musei Nationalis Pragae (Acta. Ent. Mus. Natl. Pragae) (Acta. Ent. Mus. Natl. Pragae) 58 (2) on page 416, DOI: 10.2478/aemnp-2018-0032, http://zenodo.org/record/4504894, {"references":["SHARP D. 1881: Note sur l'Ateuchus tmolus Fisch. Avec description d'une espece nouvelle du genre Synapsis. Annales de la Societe Entomologique de Belgique 25: xci-xcii.","ZIDEK J. & POKORNY S. 2010: Review of Synapsis Bates (Scarabaeidae: Scarabaeinae: Coprini), with description of a new species. Insecta Mundi 142: 1 - 21."]}

Synapsis horaki Zídek & Pokorný, 2010 (Figs 2B,H,I, 4 A–G) Synapsis horaki Zídek & Pokorný, 2010: 18, figs 12–15 (original description). Type locality. Vietnam, Vinh Phuc Province, Tam Dao, 900 m a.s.l. Type material examined. HOLOTYPE: ♁, ‘ 6–10.v.1990 | Tam Dao | Vinh Phu Distr. | Vietnam | 900 m | Jan Horák leg.’ (NMPC). Additional material examined. VIETNAM: CAO BANG PROVINCE: Pia Oac Nature Reserve, primary forest, baited pitfall trap, 5.–20.v.2016, 22°34′3.6″N, 105°53′3.3″E, 1223 m, 1 ♁, 22°34′1.4″N, 105°53′3.3″E, 1220 m, 1 ♁, 22°34′3.1″N, 105°53′4.7″E, 1220 m, 1 ♁, 22°34′3.1″N 105°53′4.4″E, 1213 m, 1 ♀, Bùi Văn Bắc leg. (all in VNUF). Diagnosis. Hypomeral cavities present; surface sparsely punctate, and not covered by macrosetae. Mesepisternal surface flat and rugose. Genae unexpanded. Frons unarmed. Pronotal anterolateral angles not protruding. Elytral striae weak and indistinctly punctate; elytral interval 2 swollen near base. Ventral surface of femora densely punctate. Description. Body length 17.5–20.1 mm, body width 10.8–12.6 mm. Colour: Dorsal surface black and glabrous. Ventral surface black on head and shiny black on thorax, abdomen and femora. Reddish brown macrosetae upon legs and pronotal margins. Mouthparts, maxillary palpi and tarsi reddish brown. Antennae brown; antennomeres IV–VI darker than other antennomeres. Head nearly semicircular, 4.4–5.3 mm long, and 7.5–8.5 mm wide. Clypeal surface extremely rugose; apex strongly and deeply emarginated, V-shaped; distance between apices of clypeal denticles (DDC) 1.4–1.6 mm; anterior margin flexed upwards with few reddish setae. Genae rectangular, quite distinctly separated from clypeus and frons by well-defined suture; surface strongly rugose and weakly punctate; margins of anterolateral angles with dense reddish macrosetae. Frons unarmed, only slightly swollen; surface weakly rugose and punctate. Antennae with 9 antennomeres; length of antennomere I approx. 1.4 mm, equal in length to antennomeres II–VI combined; antennal club approx. 1.4 mm. Prothorax. Pronotum transverse, 4.8–5.1 mm long, and 9.3–10.6 mm wide, widest at anterior quarter; pronotal disc almost indistinctly punctate, except for small weakly punctured areas near base and sides (at 30× magnification); anterolateral angles sharp and not protruding. Two lateral carinae on each side of pronotum clearly distinct; margin of outer carina with dense reddish brown macrosetae; area between carinae smooth. Hypomeral cavities present; surface of cavities weakly and sparsely punctate, and without macrosetae. Meso-metaventrum plate almost smooth, with posterior median weak groove, and with distinct excavation near metacoxae; surface of sides and anterior part sparsely and weakly punctate. Pterothorax. Elytra 10.4–12.1 mm long, 10.8–12.6 mm wide, with weak and indistinctly punctured striae. Elytral intervals convex, smooth and impunctate (at 30× magnification); interval 2 swollen near base. Mesepimeron and metepisternum flat, granulose and without macrosetae. Legs. Ventral surface of profemora strongly, coarsely and quite equally punctate; macrosetae upon profemoral margin reddish brown and long, denser in anterior margin. Protibia (ProTiL 3.2–3.6 mm, ProTiW 2.3–2.9 mm) with three broad and flat lateral teeth; protibial spurs (ProTiSL 1.4–1.6 mm) sharp, strongly curved outwards near apex, and equal in length to protibial tarsus. Ventral surface of mesofemora strongly and unequally punctate; punctures becoming denser on third posteior part. Mesotibia (MesoTiL 3.5–4 mm, MesoTiW 1.3–1.5 mm) with two sharp spurs (1 stMesoTiSL 2.0– 2.4 mm, 2 ndMesoTiSL 1.1–1.3 mm). Ventral surface of metafemora strongly and unequally punctate; punctures denser on posterior half of metafemora. Metatibia (MetaTiL 4.7–5.1 mm, MetaTiW 1.3–1.5 mm) elongate and slightly curved. Metatarsus length 3.9–4.1 mm, with 5 metatarsomeres nearly similar in size. Table 2 (on this and the opposite page). Morphological comparsions between the new species and its congeners compiled after GILLET (1911), ARROW (1931), BALTHASAR (1963), MASUMOTO (1973, 1996), HANBOONSONG & MASUMOTO (1999), KRÁL & REJSEK (2000), KRÁL (2002), OCHI & KON (2007), OCHI et al. (2008), ZÍDEK & POKORNÝ (2010). Abdomen and pygidium. Abdominal ventrites opaque, indistinctly punctate, and narrower at midline. Pygidium 2.2–2.5 mm long, 4.0– 4.8 mm wide; surface slightly convex, scabrous, and with mixture of punctures and rugosities. Aedeagus. Phallobase length 3.4–3.6 mm (in lateral view); basal suture with strong swelling at middle. Parameres length 2.0– 2.2 mm (in lateral view). Phallobase and parameres forming angle> 130 o. Sexual dimorphism. Based on an examination of the four specimens (3 males and 1 female), we did not find significant differences in morphological characters between both sexes, except for the colour of compound eyes, being black in the female but yellow in males (Figs 2H, I). This finding is consistent with the observed sexes of S. puluongensis sp. nov., raising the possibility of using this character to distinguish both sexes of these two species. Biology. All four specimens were collected in the Pia Oac Nature Reserve. The habitat is primary forests at an elevation of 1220 m a.s.l. characterized by a forest canopy cover ranging from 76 to 95%. The percentage of exposed soil was 0–5%, with 6–25% herbaceous plant layer and leaf litter cover of 96–100%. The forests has a complex structure with various storeys. Dominant trees range from 20 to 30 m tall and belong mainly to two dominant families: Fagaceae (Castanopsis spp., Lithocarpus spp., Castanea spp.) and Lauraceae (Litsea spp., Cinnamomum spp., Machilus spp.), the herbaceous and parasitic plants comprised Poaceae, Asteraceae, Orchidaceae and Loranthaceae. Remarks. So far, Synapsis horaki was known only from the holotype specimen collected in the Tam Dao National Park, Vinh Phuc Province, northern Vietnam. The herein presented specimens constitute a new record for the Cao Bang Province and the first known female. Morphometric measurements are summarized in the Table 1., Published as part of Bui, Van Bac & Bonkowski, Michael, 2018, Synapsis puluongensis sp. nov. and redescription of S. horaki (Coleoptera: Scarabaeidae), with a key to Vietnamese species, pp. 407-418 in Acta Entomologica Musei Nationalis Pragae (Acta. Ent. Mus. Natl. Pragae) (Acta. Ent. Mus. Natl. Pragae) 58 (2) on pages 413-416, DOI: 10.2478/aemnp-2018-0032, http://zenodo.org/record/4504894, {"references":["ZIDEK J. & POKORNY S. 2010: Review of Synapsis Bates (Scarabaeidae: Scarabaeinae: Coprini), with description of a new species. Insecta Mundi 142: 1 - 21.","GILLET J. J. E. 1911: Coprides nouveaux de la region orientale et remarques synonymiques. Annales de la Societe Entomologique de Belgique 55: 313 - 314.","ARROW G. J. 1931: The fauna of British India, including Ceylon and Burma. Coleoptera Lamellicornia. Part III (Coprinae). Taylor and Francis, London, 428 pp.","BALTHASAR V. 1963: Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region. Coleoptera: Lamellicornia. Band 1. Allgemeiner Teil, Systematischer Teil: 1. Scarabaeinae, 2. Coprinae (Pinotini, Coprini). Verlag der Tschechoslowakischen Akademie der Wissenschaften, Prag, 391 pp.","MASUMOTO K. 1973: Observation of the nidification of Synapsis davidi Fairmaire. Entomological Review of Japan 25: 60 - 63.","MASUMOTO K. 1996: Coprophagid-beetles from Northwest Thailand (X) (Coleoptera, Scarabaeidae). Entomological Review of Japan 50: 87 - 94.","HANBOONSONG Y. & MASUMOTO K. 1999: Dung beetles of Thailand, Part 1. Genus Synapsis. Elytra 27: 453 - 462.","KRAL D. & REJSEK J. 2000: Synapsis naxiorum sp. n. from Yunnan (Coleoptera: Scarabaeidae). Acta Societatis Zoologicae Bohemicae 64: 267 - 270.","KRAL D. 2002: Distribution and taxonomy of some Synapsis species, with description of S. strnadi sp. n. from Vietnam (Coleoptera: Scarabaeidae). Acta Societatis Zoologicae Bohemicae 66: 279 - 289.","OCHI T. & KON M. 2007: A new species of the genus Synapsis (Coleoptera, Scarabaeidae) from Laos. Elytra 35: 91 - 95.","OCHI T., KON M. & KAWAHARA M. 2008: Notes on the coprophagous scarabaeid beetles (Coleoptera: Scarabaeidae) from Southeast Asia (XIX). Three new taxa of Synapsis from Southeast Asia. Entomological Review of Japan 63: 191 - 199.","SHARP D. 1881: Note sur l'Ateuchus tmolus Fisch. Avec description d'une espece nouvelle du genre Synapsis. Annales de la Societe Entomologique de Belgique 25: xci-xcii."]}

Key to species of Synapsis recorded from Vietnam The key is based on the specimens examined by us in NMPC and VNUF as well as on literature data (ARROW 1931, BALTHASAR 1963, KRÁL & REJSEK 2000, KRÁL 2002, HANBOONSONG & MASUMOTO 1999, OCHI & KON 2007, OCHI et al. 2008, ZÍDEK & POKORNÝ 2010). 1(6) Hypomeral cavities present (Figs 2 E–F). 2(3) Elytral interval 2 near base not swollen (Fig. 2A)........................................... S. puluongensis sp. nov. 3(2) Elytral interval 2 near base swollen (Fig. 2B). 4(5) Metafemora in ventral view densely punctate, body length 17.5–24.0 mm................................................................... S. horaki Zídek & Pokorný, 2010 5(4) Metafemora in ventral view not punctate, body length 27.0–29.0 mm (Fig. 2D).................................................................................. S. yama Gillet, 1911 6(7) Hypomeral cavities absent. 7(10) Frons with minor horn or medial tubercle; mesepisternal cavities absent. 8(9) Genae not expanded, anterolateral angle of pronotum not dentate (Fig. 5A)....................................................................................... S. simplex Sharp, 1875 9(8) Genae expanded; anterolateral angle of pronotum tridentate (Fig. 5B)............. S. tridens Sharp, 1881 10(11) Frons without minor horn or only with medial tubercle; mesepisternal cavities present and covered with red setae (Figs 6 A–C, F). 11(12) Anterolateral angles of pronotum nearly rectangular (90°). Lateral angles of genae obtuse, rounded. Metafemora in ventral view sparsely punctate (Figs 6A, D)................. S. ovalis Boucomont, 1920 12(11) Anterolateral angles of pronotum about 135°. Lateral angles of genae rather sharp, metafemora in ventral view densely punctate (Figs 6B, E).............................................................. S. strnadi Král, 2002, Published as part of Bui, Van Bac & Bonkowski, Michael, 2018, Synapsis puluongensis sp. nov. and redescription of S. horaki (Coleoptera: Scarabaeidae), with a key to Vietnamese species, pp. 407-418 in Acta Entomologica Musei Nationalis Pragae (Acta. Ent. Mus. Natl. Pragae) (Acta. Ent. Mus. Natl. Pragae) 58 (2) on pages 416-417, DOI: 10.2478/aemnp-2018-0032, http://zenodo.org/record/4504894, {"references":["ARROW G. J. 1931: The fauna of British India, including Ceylon and Burma. Coleoptera Lamellicornia. Part III (Coprinae). Taylor and Francis, London, 428 pp.","BALTHASAR V. 1963: Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region. Coleoptera: Lamellicornia. Band 1. Allgemeiner Teil, Systematischer Teil: 1. Scarabaeinae, 2. Coprinae (Pinotini, Coprini). Verlag der Tschechoslowakischen Akademie der Wissenschaften, Prag, 391 pp.","KRAL D. & REJSEK J. 2000: Synapsis naxiorum sp. n. from Yunnan (Coleoptera: Scarabaeidae). Acta Societatis Zoologicae Bohemicae 64: 267 - 270.","KRAL D. 2002: Distribution and taxonomy of some Synapsis species, with description of S. strnadi sp. n. from Vietnam (Coleoptera: Scarabaeidae). Acta Societatis Zoologicae Bohemicae 66: 279 - 289.","HANBOONSONG Y. & MASUMOTO K. 1999: Dung beetles of Thailand, Part 1. Genus Synapsis. Elytra 27: 453 - 462.","OCHI T. & KON M. 2007: A new species of the genus Synapsis (Coleoptera, Scarabaeidae) from Laos. Elytra 35: 91 - 95.","OCHI T., KON M. & KAWAHARA M. 2008: Notes on the coprophagous scarabaeid beetles (Coleoptera: Scarabaeidae) from Southeast Asia (XIX). Three new taxa of Synapsis from Southeast Asia. Entomological Review of Japan 63: 191 - 199.","ZIDEK J. & POKORNY S. 2010: Review of Synapsis Bates (Scarabaeidae: Scarabaeinae: Coprini), with description of a new species. Insecta Mundi 142: 1 - 21.","GILLET J. J. E. 1911: Coprides nouveaux de la region orientale et remarques synonymiques. Annales de la Societe Entomologique de Belgique 55: 313 - 314.","SHARP D. 1881: Note sur l'Ateuchus tmolus Fisch. Avec description d'une espece nouvelle du genre Synapsis. Annales de la Societe Entomologique de Belgique 25: xci-xcii."]}

Synapsis horaki Zídek & Pokorný, 2010 (Figs 2B,H,I, 4 A–G) Synapsis horaki Zídek & Pokorný, 2010: 18, figs 12–15 (original description). Type locality. Vietnam, Vinh Phuc Province, Tam Dao, 900 m a.s.l. Type material examined. HOLOTYPE: ♁, ‘ 6–10.v.1990 | Tam Dao | Vinh Phu Distr. | Vietnam | 900 m | Jan Horák leg.’ (NMPC). Additional material examined. VIETNAM: CAO BANG PROVINCE: Pia Oac Nature Reserve, primary forest, baited pitfall trap, 5.–20.v.2016, 22°34′3.6″N, 105°53′3.3″E, 1223 m, 1 ♁, 22°34′1.4″N, 105°53′3.3″E, 1220 m, 1 ♁, 22°34′3.1″N, 105°53′4.7″E, 1220 m, 1 ♁, 22°34′3.1″N 105°53′4.4″E, 1213 m, 1 ♀, Bùi Văn Bắc leg. (all in VNUF). Diagnosis. Hypomeral cavities present; surface sparsely punctate, and not covered by macrosetae. Mesepisternal surface flat and rugose. Genae unexpanded. Frons unarmed. Pronotal anterolateral angles not protruding. Elytral striae weak and indistinctly punctate; elytral interval 2 swollen near base. Ventral surface of femora densely punctate. Description. Body length 17.5–20.1 mm, body width 10.8–12.6 mm. Colour: Dorsal surface black and glabrous. Ventral surface black on head and shiny black on thorax, abdomen and femora. Reddish brown macrosetae upon legs and pronotal margins. Mouthparts, maxillary palpi and tarsi reddish brown. Antennae brown; antennomeres IV–VI darker than other antennomeres. Head nearly semicircular, 4.4–5.3 mm long, and 7.5–8.5 mm wide. Clypeal surface extremely rugose; apex strongly and deeply emarginated, V-shaped; distance between apices of clypeal denticles (DDC) 1.4–1.6 mm; anterior margin flexed upwards with few reddish setae. Genae rectangular, quite distinctly separated from clypeus and frons by well-defined suture; surface strongly rugose and weakly punctate; margins of anterolateral angles with dense reddish macrosetae. Frons unarmed, only slightly swollen; surface weakly rugose and punctate. Antennae with 9 antennomeres; length of antennomere I approx. 1.4 mm, equal in length to antennomeres II–VI combined; antennal club approx. 1.4 mm. Prothorax. Pronotum transverse, 4.8–5.1 mm long, and 9.3–10.6 mm wide, widest at anterior quarter; pronotal disc almost indistinctly punctate, except for small weakly punctured areas near base and sides (at 30× magnification); anterolateral angles sharp and not protruding. Two lateral carinae on each side of pronotum clearly distinct; margin of outer carina with dense reddish brown macrosetae; area between carinae smooth. Hypomeral cavities present; surface of cavities weakly and sparsely punctate, and without macrosetae. Meso-metaventrum plate almost smooth, with posterior median weak groove, and with distinct excavation near metacoxae; surface of sides and anterior part sparsely and weakly punctate. Pterothorax. Elytra 10.4–12.1 mm long, 10.8–12.6 mm wide, with weak and indistinctly punctured striae. Elytral intervals convex, smooth and impunctate (at 30× magnification); interval 2 swollen near base. Mesepimeron and metepisternum flat, granulose and without macrosetae. Legs. Ventral surface of profemora strongly, coarsely and quite equally punctate; macrosetae upon profemoral margin reddish brown and long, denser in anterior margin. Protibia (ProTiL 3.2–3.6 mm, ProTiW 2.3–2.9 mm) with three broad and flat lateral teeth; protibial spurs (ProTiSL 1.4–1.6 mm) sharp, strongly curved outwards near apex, and equal in length to protibial tarsus. Ventral surface of mesofemora strongly and unequally punctate; punctures becoming denser on third posteior part. Mesotibia (MesoTiL 3.5–4 mm, MesoTiW 1.3–1.5 mm) with two sharp spurs (1 stMesoTiSL 2.0– 2.4 mm, 2 ndMesoTiSL 1.1–1.3 mm). Ventral surface of metafemora strongly and unequally punctate; punctures denser on posterior half of metafemora. Metatibia (MetaTiL 4.7–5.1 mm, MetaTiW 1.3–1.5 mm) elongate and slightly curved. Metatarsus length 3.9–4.1 mm, with 5 metatarsomeres nearly similar in size. Table 2 (on this and the opposite page). Morphological comparsions between the new species and its congeners compiled after GILLET (1911), ARROW (1931), BALTHASAR (1963), MASUMOTO (1973, 1996), HANBOONSONG & MASUMOTO (1999), KRÁL & REJSEK (2000), KRÁL (2002), OCHI & KON (2007), OCHI et al. (2008), ZÍDEK & POKORNÝ (2010). Abdomen and pygidium. Abdominal ventrites opaque, indistinctly punctate, and narrower at midline. Pygidium 2.2–2.5 mm long, 4.0– 4.8 mm wide; surface slightly convex, scabrous, and with mixture of punctures and rugosities. Aedeagus. Phallobase length 3.4–3.6 mm (in lateral view); basal suture with strong swelling at middle. Parameres length 2.0– 2.2 mm (in lateral view). Phallobase and parameres forming angle> 130 o. Sexual dimorphism. Based on an examination of the four specimens (3 males and 1 female), we did not find significant differences in morphological characters between both sexes, except for the colour of compound eyes, being black in the female but yellow in males (Figs 2H, I). This finding is consistent with the observed sexes of S. puluongensis sp. nov., raising the possibility of using this character to distinguish both sexes of these two species. Biology. All four specimens were collected in the Pia Oac Nature Reserve. The habitat is primary forests at an elevation of 1220 m a.s.l. characterized by a forest canopy cover ranging from 76 to 95%. The percentage of exposed soil was 0–5%, with 6–25% herbaceous plant layer and leaf litter cover of 96–100%. The forests has a complex structure with various storeys. Dominant trees range from 20 to 30 m tall and belong mainly to two dominant families: Fagaceae (Castanopsis spp., Lithocarpus spp., Castanea spp.) and Lauraceae (Litsea spp., Cinnamomum spp., Machilus spp.), the herbaceous and parasitic plants comprised Poaceae, Asteraceae, Orchidaceae and Loranthaceae. Remarks. So far, Synapsis horaki was known only from the holotype specimen collected in the Tam Dao National Park, Vinh Phuc Province, northern Vietnam. The herein presented specimens constitute a new record for the Cao Bang Province and the first known female. Morphometric measurements are summarized in the Table 1.

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

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

Aporcelinus paramamillatus sp. n. (Figs 1���2) Material examined. Six females from one location, in good condition. Morphometrics. See Table 1. Description. Female: Moderately slender (a = 21���26) nematodes of medium size, 1.71���2.14 mm long. Habitus visibly curved ventrad upon fixation, C-shaped. Cuticle two-layered, 4 ��m thick at anterior region and mid-body and 4���7 ��m on tail; inner layer thicker and more refractive than the outer layer. Lateral chords 10���14 ��m broad or 12���19 % of mid-body diameter. Body pores often obscure, but two cervical dorsal and two ventral are sometimes distinct at the level of the odontophore. Lip region somewhat angular, offset by weak constriction, 2.6���2.8 times as wide as high and 26���31 % of body diameter at neck base; lips moderately separated, with protruding labial and cephalic papillae. Amphid fovea cup-shaped, its opening 10���12 ��m broad or about one-half of lip region diameter. Cheilostom nearly as long as wide, with no specialization. Odontostyle strong, slightly (1.1���1.2 times) longer than lip region diameter, 5.2���6.5 times as long as wide and 1.1���1.5 % of total body length; aperture 12 ��m or 46���50 % of its total length. Guiding ring simple but distinct and somewhat plicate, located at 12���13 ��m or 0.5 times the lip region diameter from the anterior end. Odontophore rod-like, 1.7 ���2.0 times the odontostyle in length. Pharynx consisting of a slender but muscular anterior section enlarging very gradually into the posterior expansion that is 6.4���8.3 times as long as wide, 2.7���3.4 times the corresponding body diameter long, and occupies 47���56 % of total neck length; gland nuclei located as follows (n = 1): DO = 58, DN = 62, S 1 N 1 = 75, S 1 N 2 = 79, S 2 N = 92. Nerve ring at 163���212 ��m or 31���35 % of total neck length from the anterior end. Pharyngointestinal junction bearing a distinctly developed dorsal lobe; cardia conical, 12��� 18 x 8���15 ��m. Genital system didelphic-amphidelphic, with both branches almost equally and moderately developed, 157���224 ��m long or 9���12 % of total body length: ovaries 57���177 ��m long, surpassing the oviduct-uterus junction; oviduct 65���106 ��m long or 0.9���1.4 times the body diameter, and consisting of a slender portion and a moderately developed pars dilatata; a narrowing separates oviduct and uterus; uterus 71���94 ��m long or 1.0��� 1.6 times the corresponding body diameter long, tripartite, consisting of a short and wider proximal region, a narrower intermediate section with no lumen, and a nearly spherical distal part; uterine egg 90 x 55 ��m, with irregular shell; vagina extending inwards 26���32 ��m or 32���43 % of body diameter, with pars proximalis 16��� 22 x 18���22 ��m and somewhat sigmoid walls surrounded by weak musculature, pars refringens consisting of two drop-shaped pieces 6��� 7 x 4���6 ��m and a combined width of 9���12 ��m, and pars distalis 3���5 ��m long and visibly refractive; vulva a transverse slit. Prerectum 1.8���2.2, rectum 1.0��� 1.2 times the anal body diameter long. Tail conical with finely rounded tip, dorsally and ventrally convex, ending in a short peg-like terminus, occasionally slightly re-curved dorsad; hyaline portion very short; caudal pores two pairs at the anterior part of tail, one subdorsal, the other sublateral. Male: Unknown. Diagnosis. The new species is characterized by its 1.71���2.14 mm long body, lip region offset by weak constriction and 21���22 ��m broad, odontostyle 22���24 ��m at its ventral side or 1.1���1.2 times the lip region diameter, neck 451���503 ��m long, pharyngeal expansion 183���251 ��m long or 46���56 % of total neck length, uterus tripartite and 71���94 ��m long or 1.0��� 1.6 times the corresponding body diameter long, V = 53���54, tail conical (27���34 ��m, c = 50���77, c��� = 0.6���0.9) ending in a peg-like terminal portion and not visibly re-curved dorsad, and male absent. Relationships. This species is very similar to A. mamillatus (Williams, 1959) ��lvarez-Ortega & Pe��a- Santiago, 2011 in its general morphology and many morphometrics, but it can be distinguished by several significant differences: broader lip region (21���22 vs 19���20 ��m), smaller odontostyle (22���24 vs 25���29 ��m, 1.1���1.2 vs 1.3���1.5 times the lip region diameter, n= 9) with shorter aperture (46���50 vs 57���61 % of its total length), more posterior S 2 N (92 vs 83���86) and more posterior vulva (V = 53���54 vs V = 49���52). Type locality and habitat. Northern Vietnam, Cat Ba National Park, Hai Phong Province (GPS coordinates: 20 �� 47 ' 45 '' N and 107 �� 00' 40 ''E, elevation 170 m), in soil of a secondary forest with Cinnamomum sp. as the dominant plant species. One female was collected near the same location in the rhizosphere of Zea mays L. growing in an area of intensive agricultural crops alongside the National Park. Type material. Female holotype and four females paratypes deposited in the nematode collection of the University of Ja��n, Spain. One female paratype in the nematode collection of the Institute of Ecology and Biological Resources (IEBR), Hanoi, Vietnam. Etymology. The specific epithet means ���near mamillatus ��� as the new species resembles A. mamillatus in many aspects., Published as part of Nguyen, Thi Anh Duong, Abolafia, Joaqu��n, Bonkowski, Michael & Santiago, Reyes Pe��a-, 2016, Two new species of the genus Aporcelinus Andr��ssy, 2009 (Nematoda, Dorylaimida, Aporcelaimidae) from Vietnam, pp. 550-560 in Zootaxa 4103 (6) on pages 551-554, DOI: 10.11646/zootaxa.4103.6.5, http://zenodo.org/record/271219, {"references":["Williams, J. R. (1959) Studies on the nematode soil fauna of sugarcane fields in Mauritius. 3. Dorylaimidae (Dorylaimoidea, Enoplida). Mauritius Sugar Industry Research Institute Occasional Paper, 3, 1 - 28."]}

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

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

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

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

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

Nguyen, Thi Anh Duong, Bonkowski, Michael, Peña-Santiago, Reyes, Abolafia, Joaquín (2016): Re-description of Cephalobus topali Andrássy, 1970 (Rhabditida, Cephalobidae) from Vietnam, and transfer to Acrobeloides (Cobb, 1924) Thorne, 1937. Zootaxa 4092 (4): 593-600, DOI: http://doi.org/10.11646/zootaxa.4092.4.9

Aporcelinus paraseychellensis sp. n. (Figs 3 ���5) Syn. Aporcelinus seychellensis apud Vinciguerra et al. (2014), nec Andr��ssy (2009) Material examined. Three females and one male from two locations, in acceptable condition. Morphometrics. See Table 2. Locality Cat Ba National Park Huu Lien Natural Reserve (Hai Phong Province) (Lang Son Province) Habitat Cinnamomum sp Machilus sp. and Dimocarpus Holotype Paratypes sp. Description. Adult: Stout (a = 16���20) nematodes of medium size, 1.20���1.46 mm long. Habitus variably curved ventrad upon fixation, C-shaped in female, G-shaped in male. Cuticle two-layered, 3���4 ��m thick at anterior region, 3.5 ���5.0 ��m in mid-body and 4.0��� 6.5 ��m on tail; inner layer thicker and more refractive than the outer layer. Lateral chord 9 ��m broad or 10���13 % of mid-body diameter. Body pores often obscure, but two cervical pores are visible on both dorsal and ventral sides at level of odontostyle plus odontophore. Lip region angular, offset by constriction; lips high, separated, with protruding labial and cephalic papillae. Amphid fovea cup-shaped, its opening 9���11 ��m broad or 53���63 % of lip region diameter. Cheilostom nearly as long as wide, with no specialization. Odontostyle strong, 1.1���1.4 times longer than lip region diameter, 6.2���7.7 times as long as wide and 1.4���1.9 % of total body length; aperture 10���11 ��m or 46���48 % of its total length. Guiding ring simple but distinct, somewhat plicate, located at 9���10 ��m or 0.5���0.6 times the lip region diameter from the anterior end. Odontophore rod-like, 1.5���1.8 times the odontostyle length. Pharynx consisting of a slender but muscular anterior section enlarging very gradually in the posterior expansion, which is 5.4���6.8 times as long as wide, 2.8���3.3 times the corresponding body diameter and occupies 49���51 % of total neck length; gland nuclei located as follows (n = 1): DO = 58, DN = 62, S 1 N 1 = 75, S 1 N 2 = 82, S 2 N = 90. Nerve ring at 113 ��m or 31 % of total neck length from the anterior end. Pharyngo-intestinal junction bearing a distinctly developed dorsal lobe; cardia conical, 10��� 14 x 11���12 ��m. A dorsal cell mass is present at level of the pharyngo-intestinal junction. Female: Genital system didelphic-amphidelphic, with both branches equally and moderately developed, 176���308 ��m long or 17���21 % of total body length: ovaries very variably sized, 79���236 ��m long; oviduct 55���94 ��m long or 0.8���1.1 times the body diameter and consisting of a slender portion made of prismatic cells and a moderately developed pars dilatata with perceptible lumen; a narrowing separates oviduct and uterus, but a distinct sphincter is lacking; uterus 176���241 ��m long or 2.4���2.7 times the corresponding body diameter, tripartite, consisting of a short and wider proximal region, a longer and much narrower intermediate section with no lumen, and a nearly spherical distal part; vagina extending inwards 33���36 ��m or 40���47 % of body diameter, with pars proximalis 22��� 25 x 15���20 ��m and somewhat sigmoid walls surrounded by weak musculature, pars refringens consisting of two drop-shaped or trapezoidal pieces 7��� 8 x 4���5 ��m and a combined width of 9���10 ��m, and pars distalis 5���7 ��m long and visibly refractive; vulva a transverse slit. Prerectum 1.5 ���2.0, rectum 1.2���1.3 times the anal body diameter in length. Tail conical with acute tip, ventrally nearly straight or slightly convex, and dorsally first convex and then concave, the tail being slightly re-curved dorsad; cuticle extending until the tail tip, so that a hyaline portion is not observed; caudal pores two pairs, at the middle of tail, one subdorsal, the other lateral. Male: Prerectum 2.3, cloaca 1.7 times the body diameter at the cloacal aperture. Genital system diorchic, with opposite testes. In addition to the adcloacal pair, located at 9 ��m from the cloacal aperture, there is a series of 12 low, regularly spaced (10���16 ��m apart) ventromedian supplements, the most posterior three located within the range of spicules. Spicules dorylaimid, 4.6 times longer than wide and 1.8 times longer than body diameter at the cloacal aperture: dorsal side regularly convex and ventral side with barely perceptible hump and hollow; curvature 136 ��; head 8 x 7 ��m, with the dorsal side somewhat longer than the ventral one and weakly curved; median piece occupying less than one-third (29 %) of maximum spicule width, reaching the posterior end of the spicule and with a visibly furcate tip; posterior end of spicules 6 ��m wide. Lateral guiding pieces 17 ��m long or 6.1 times as long as wide, slightly sigmoid and tapering posteriorly. Tail straight ventrally, dorsally first convex and then very weakly concave. Diagnosis. The new species is characterized by its 1.20���1.46 mm long body, lip region offset by constriction and 16���18 ��m broad, odontostyle 20 ��m at its ventral side or 1.1���1.4 times the lip region diameter, neck 354���368 ��m long, pharyngeal expansion 175���186 ��m long or 49���51 % of total neck length, a dorsal cell mass present at the pharyngo-intestinal junction, uterus tripartite and 176���241 ��m, or 2.4���2.7 times the corresponding body diameter, long, V = 52���54, tail conical with acute tip and slightly recurved dorsad (33���42 ��m, c = 33���45, c��� = 0.9���1.2), spicules 62 ��m long, and 12 regularly spaced ventromedian supplements, three of them within the range of spicules. Relationships. The new species is very similar to A. seychellensis Andr��ssy, 2011 in its general morphology and morphometrics, but differs from the type population of that species in its larger general size (body length 1.20���1.46 (n = 4) vs 0.92���1.07 mm (n= 4); neck length 354���368 vs 272���330 ��m), much larger genital branches (17���21 % vs 7���13 % of total body length each), more posterior vulva (V = 52���54 vs V = 45���52), shorter female tail (32���36 vs 36���40 ��m, c = 33���45 vs c = 25���29) and male present (vs absent). Especially relevant in the type population are the very short female genital branches, probably because the uterus is short and simple. Vinciguerra et al. (2014) studied two A. seychellensis populations from Ecuador that differ from the type populations in their larger general size [body 1.08���1.55 (n= 26) vs 0.92���1.07 (n = 4) mm long] and comparatively shorter female tail (c = 29���42 vs c = 25���29). Besides, the female genital branches are larger (12���22 vs 7���13 % of total body length), but lower values in the Ecuador material may be due to the convoluted, tripartite uterus. Leaving aside the fewer ventromedian supplements (7���9 vs 12; two vs three situated within the range of spicules) in males from Ecuador, there is no significant difference between this material and the Vietnamese specimens herein studied. Taking into consideration that only one male of the new species has been collected, the differences in number and arrangement of ventromedian supplements are provisionally regarded as intraspecific variability and, consequently, we consider the specimens from Ecuador and Vietnam to be conspecific. Type locality and habitat. Northern Vietnam, Cat Ba National Park, Hai Phong Province (GPS coordinates: 20 �� 47 ' 45 '' N and 107 �� 00' 40 ''E, elevation 170 m), collected from soil in a tropical forest with Cinnamomum sp. as dominant plant species. Other locality and habitat. Northern Vietnam, Huu Lien Natural Reserve, Lang Son Province, (GPS coordinates: 21 �� 42 ' 51 '' N and 106 �� 21 ' 48 ''E, elevation 502 m), in soil of the forest where the dominant plant species were Machilus sp. and Dimocarpus sp. Type material. Female holotype, female paratype and male paratype deposited in the nematode collection of the University of Ja��n, Spain. One female paratype deposited in the nematode collection of the Institute of Ecology and Biological Resources (IEBR), Hanoi, Vietnam. Etymology. The specific epithet means ���near seychellensis ��� as the new species resembles A. seychellensis in many aspects., Published as part of Nguyen, Thi Anh Duong, Abolafia, Joaqu��n, Bonkowski, Michael & Santiago, Reyes Pe��a-, 2016, Two new species of the genus Aporcelinus Andr��ssy, 2009 (Nematoda, Dorylaimida, Aporcelaimidae) from Vietnam, pp. 550-560 in Zootaxa 4103 (6) on pages 555-559, DOI: 10.11646/zootaxa.4103.6.5, http://zenodo.org/record/271219, {"references":["Vinciguerra, M. T., Orselli, L. & Clausi, M. (2014) One new and two known species of Aporcelinus Andrassy, 2009 and a new species of Coomansinema Ahmad & Jairajpuri, 1989 (Nematoda: Dorylaimida). Nematology, 16, 303 - 322. http: // dx. doi. org / 10.1163 / 15685411 - 00002767","Andrassy, I. (2011) Phylum Nematoda. In: Gerlach, J. (Ed.), Crustacea, Platyhelminthes, Nematoda, Annelida and Tardigrada of the Seychelles Islands: 15 - 57. Siri Scientific Press, Manchester. 160 pp."]}

Luu, Vinh Quang, Nguyen, Truong Quang, Le, Minh Duc, Bonkowski, Michael, Ziegler, Thomas (2016): A new species of karst-dwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa 4079 (1): 87-102, DOI: http://doi.org/10.11646/zootaxa.4079.1.6

Aporcelinus paraseychellensis sp. n. (Figs 3 ‒5) Syn. Aporcelinus seychellensis apud Vinciguerra et al. (2014), nec Andrássy (2009) Material examined. Three females and one male from two locations, in acceptable condition. Morphometrics. See Table 2. Locality Cat Ba National Park Huu Lien Natural Reserve (Hai Phong Province) (Lang Son Province) Habitat Cinnamomum sp Machilus sp. and Dimocarpus Holotype Paratypes sp. Description. Adult: Stout (a = 16–20) nematodes of medium size, 1.20–1.46 mm long. Habitus variably curved ventrad upon fixation, C-shaped in female, G-shaped in male. Cuticle two-layered, 3–4 µm thick at anterior region, 3.5 –5.0 µm in mid-body and 4.0– 6.5 µm on tail; inner layer thicker and more refractive than the outer layer. Lateral chord 9 µm broad or 10–13 % of mid-body diameter. Body pores often obscure, but two cervical pores are visible on both dorsal and ventral sides at level of odontostyle plus odontophore. Lip region angular, offset by constriction; lips high, separated, with protruding labial and cephalic papillae. Amphid fovea cup-shaped, its opening 9–11 µm broad or 53–63 % of lip region diameter. Cheilostom nearly as long as wide, with no specialization. Odontostyle strong, 1.1–1.4 times longer than lip region diameter, 6.2–7.7 times as long as wide and 1.4–1.9 % of total body length; aperture 10–11 µm or 46–48 % of its total length. Guiding ring simple but distinct, somewhat plicate, located at 9–10 µm or 0.5–0.6 times the lip region diameter from the anterior end. Odontophore rod-like, 1.5–1.8 times the odontostyle length. Pharynx consisting of a slender but muscular anterior section enlarging very gradually in the posterior expansion, which is 5.4–6.8 times as long as wide, 2.8–3.3 times the corresponding body diameter and occupies 49–51 % of total neck length; gland nuclei located as follows (n = 1): DO = 58, DN = 62, S 1 N 1 = 75, S 1 N 2 = 82, S 2 N = 90. Nerve ring at 113 µm or 31 % of total neck length from the anterior end. Pharyngo-intestinal junction bearing a distinctly developed dorsal lobe; cardia conical, 10– 14 x 11–12 µm. A dorsal cell mass is present at level of the pharyngo-intestinal junction. Female: Genital system didelphic-amphidelphic, with both branches equally and moderately developed, 176–308 µm long or 17–21 % of total body length: ovaries very variably sized, 79–236 µm long; oviduct 55–94 µm long or 0.8–1.1 times the body diameter and consisting of a slender portion made of prismatic cells and a moderately developed pars dilatata with perceptible lumen; a narrowing separates oviduct and uterus, but a distinct sphincter is lacking; uterus 176–241 µm long or 2.4–2.7 times the corresponding body diameter, tripartite, consisting of a short and wider proximal region, a longer and much narrower intermediate section with no lumen, and a nearly spherical distal part; vagina extending inwards 33–36 µm or 40–47 % of body diameter, with pars proximalis 22– 25 x 15–20 µm and somewhat sigmoid walls surrounded by weak musculature, pars refringens consisting of two drop-shaped or trapezoidal pieces 7– 8 x 4–5 µm and a combined width of 9–10 µm, and pars distalis 5–7 µm long and visibly refractive; vulva a transverse slit. Prerectum 1.5 –2.0, rectum 1.2–1.3 times the anal body diameter in length. Tail conical with acute tip, ventrally nearly straight or slightly convex, and dorsally first convex and then concave, the tail being slightly re-curved dorsad; cuticle extending until the tail tip, so that a hyaline portion is not observed; caudal pores two pairs, at the middle of tail, one subdorsal, the other lateral. Male: Prerectum 2.3, cloaca 1.7 times the body diameter at the cloacal aperture. Genital system diorchic, with opposite testes. In addition to the adcloacal pair, located at 9 µm from the cloacal aperture, there is a series of 12 low, regularly spaced (10–16 µm apart) ventromedian supplements, the most posterior three located within the range of spicules. Spicules dorylaimid, 4.6 times longer than wide and 1.8 times longer than body diameter at the cloacal aperture: dorsal side regularly convex and ventral side with barely perceptible hump and hollow; curvature 136 º; head 8 x 7 µm, with the dorsal side somewhat longer than the ventral one and weakly curved; median piece occupying less than one-third (29 %) of maximum spicule width, reaching the posterior end of the spicule and with a visibly furcate tip; posterior end of spicules 6 µm wide. Lateral guiding pieces 17 µm long or 6.1 times as long as wide, slightly sigmoid and tapering posteriorly. Tail straight ventrally, dorsally first convex and then very weakly concave. Diagnosis. The new species is characterized by its 1.20–1.46 mm long body, lip region offset by constriction and 16–18 µm broad, odontostyle 20 µm at its ventral side or 1.1–1.4 times the lip region diameter, neck 354–368 µm long, pharyngeal expansion 175–186 µm long or 49–51 % of total neck length, a dorsal cell mass present at the pharyngo-intestinal junction, uterus tripartite and 176–241 µm, or 2.4–2.7 times the corresponding body diameter, long, V = 52–54, tail conical with acute tip and slightly recurved dorsad (33–42 µm, c = 33–45, c’ = 0.9–1.2), spicules 62 µm long, and 12 regularly spaced ventromedian supplements, three of them within the range of spicules. Relationships. The new species is very similar to A. seychellensis Andrássy, 2011 in its general morphology and morphometrics, but differs from the type population of that species in its larger general size (body length 1.20–1.46 (n = 4) vs 0.92–1.07 mm (n= 4); neck length 354–368 vs 272–330 µm), much larger genital branches (17–21 % vs 7–13 % of total body length each), more posterior vulva (V = 52–54 vs V = 45–52), shorter female tail (32–36 vs 36–40 µm, c = 33–45 vs c = 25–29) and male present (vs absent). Especially relevant in the type population are the very short female genital branches, probably because the uterus is short and simple. Vinciguerra et al. (2014) studied two A. seychellensis populations from Ecuador that differ from the type populations in their larger general size [body 1.08–1.55 (n= 26) vs 0.92–1.07 (n = 4) mm long] and comparatively shorter female tail (c = 29–42 vs c = 25–29). Besides, the female genital branches are larger (12–22 vs 7–13 % of total body length), but lower values in the Ecuador material may be due to the convoluted, tripartite uterus. Leaving aside the fewer ventromedian supplements (7–9 vs 12; two vs three situated within the range of spicules) in males from Ecuador, there is no significant difference between this material and the Vietnamese specimens herein studied. Taking into consideration that only one male of the new species has been collected, the differences in number and arrangement of ventromedian supplements are provisionally regarded as intraspecific variability and, consequently, we consider the specimens from Ecuador and Vietnam to be conspecific. Type locality and habitat. Northern Vietnam, Cat Ba National Park, Hai Phong Province (GPS coordinates: 20 º 47 ' 45 '' N and 107 º 00' 40 ''E, elevation 170 m), collected from soil in a tropical forest with Cinnamomum sp. as dominant plant species. Other locality and habitat. Northern Vietnam, Huu Lien Natural Reserve, Lang Son Province, (GPS coordinates: 21 º 42 ' 51 '' N and 106 º 21 ' 48 ''E, elevation 502 m), in soil of the forest where the dominant plant species were Machilus sp. and Dimocarpus sp. Type material. Female holotype, female paratype and male paratype deposited in the nematode collection of the University of Jaén, Spain. One female paratype deposited in the nematode collection of the Institute of Ecology and Biological Resources (IEBR), Hanoi, Vietnam. Etymology. The specific epithet means ‘near seychellensis ’ as the new species resembles A. seychellensis in many aspects.

Cephalobus topali Andr��ssy, 1970 (Figs 1���3) Material examined. Six females and eight larvae from Pia Oac, Vietnam, in acceptable state of preservation. Morphometrics. See Table 1. Description. Female: Body 0.31���0.35 mm long. Habitus straight or sigmoid after fixation. Cuticle distinctly annulated, annuli 2 mm thick at mid-body. Lateral field occupying 15���24 % of mid-body diameter, with two alae (three or four longitudinal incisures, depending on either alae are slightly separate or not) arising at pharyngeal corpus level and fading away at phasmid level, appearing only one incisure posteriorly. With SEM, anterior region seen to comprise three pairs of asymmetrical conoid lips, and three low and rounded labial probolae separated from the corresponding pair of lips by an incisure; primary axils deep, U-shaped, lacking guard processes; secondary axils shallow, V-shaped, lacking guard processes. Sensillae papilliform, six labial and four cephalic. Amphids slitlike, with thickened margin. Stoma cephaloboid: cheilostom with rounded rhabdia; gymnostom narrower than cheilostom, and as wide as stegostom, the latter with small rhabdia. Pharynx also cephaloboid: slightly fusiform corpus, 2.6���3.1 times the isthmus length, with procorpus longer than metacorpus; isthmus narrower than corpus; basal bulb pyriform, bearing well developed valves at its middle. Cardia hemispheroid, surrounded by intestinal tissue. Intestine lacking specialization; its lumen often sigmoid. Nerve ring located at 56���76 % of neck (stoma + pharynx) length, at level of the posterior part of corpus, near its junction to isthmus. Excretory pore at 61���87 % of neck length, 44���48 annuli from the anterior end, at level of corpus-isthmus junction, in front of hemizonid. Deirid at 63���81 % of neck length, 49���53 annuli from the anterior end, at level of basal bulb. Reproductive system monodelphic-prodelphic: ovary posteriorly directed, with or without a double flexure posterior to vulva level; oviduct short, difficult to distinguish; spermatheca poorly developed, 0.3���0.4 times the corresponding body diameter in length; uterus tube-like, as long as the corresponding body diameter, consisting of a shorter proximal part with thin walls and distinct lumen and a longer distal section having thick walls and small lumen; post-vulval sac 0.2���0.4 times the corresponding body diameter in length; vagina extending inwards to one-third of body diameter. Rectum 1.3���1.7 anal body widths in length. Tail conoid, with 11���13 annuli on the ventral side; terminus rounded, ending in an acute or wart-shaped mucro. Phasmid situated at 29���34 % of tail length. Male: Unknown. Diagnosis (based on the 6 specimens studied). This species is characterized by its body length (306���347 ��m in females), lateral field with two alae or 3���4 longitudinal incisures, lip region with three pairs of asymmetrical conoid lips, three low and rounded labial probolae separate from the corresponding pair of lips by an incisure, Ushaped primary axils without guard processes, V-shaped secondary axils without guard processes, pharyngeal corpus slightly fusiform and 2.6���3.1 times the isthmus length, spermatheca and post-vulval sac poorly developed, female tail conoid (18���24 ��m, c = 12.7 ���17.0, c��� = 1.8���2.7) with rounded terminus ending in an acute or wartshaped mucro, and phasmid at 29���34 % of tail length. Males are unknown. Locality and habitat. Pia Oac (GPS coordinates: 22 �� 36 ' 28 ''N and 105 �� 52 ' 15 ''E, elevation 1150 m), Cao Bang Province, Vietnam, collected from the rhizosphere of Zea mays L. growing in an area of intensive agricultural crops alongside the Pia Oac Natural Reserve. * Measurements from drawings, ** from other measurements. Discussion. Cephalobus topali was originally described from only one female and three larval specimens. The material herein studied perfectly fits Andr��ssy���s (1970) description. The SEM observations reported here provide additional information that raises doubt about the position of this species. For example, instead of connecting the corresponding adjacent pairs of lips, a typical diagnostic feature of the genus Cephalobus Bastian, 1865 (see Abolafia & Pe��a-Santiago 2009), the labial probolae of the specimen examined here are clearly separated from the lips by an incisures (Fig. 3 A���C). This fits the pattern observed in representatives of the genus Acrobeloides (Cobb, 1924) Thorne, 1937 (see Abolafia & Pe��a-Santiago, 2003). Consequently, this species is transferred to the genus Acrobeloides as A. topali (Andr��ssy, 1970) comb n. Andr��ssy (1984) regarded C. topali as identical to and a junior synonym of Cephalobus quadrilineatus Eroshenko, 1968, only known to occur in Russian Far East Territories. Both species have similar morphology and morphometry (see comparison in Table 1), and both have an acute mucro at the tail tip. Later, however, Andr��ssy (2005) maintained both species as valid, but examination of specimens of the Russian species is necessary to confirm it. Provisionally, we accept that the two species are valid as differences between them were noted: isthmus size (as long as vs one-third of the corresponding body diameter), excretory pore location (at level of the corpusisthmus junction vs at isthmus posterior portion, near the basal bulb), presence of a short vs absence of a postvulval sac, and rectum length (as long as vs shorter than the corresponding body diameter). The distribution of A. topali appears to be restricted to the north of Vietnam, associated with modified soils. Andr��ssy (1970) found it among grass roots in the municipal cemetery of the district of Yen So (Hanoi), and here it is reported from further north, in agricultural plots opened in the rain forest, close to the Pia Oac Natural Reserve. In the last two years, several hundred specimens of nematodes have been examined from this area but, only a few specimens of this rare species were found, and none came from natural, unaltered areas., Published as part of Nguyen, Thi Anh Duong, Bonkowski, Michael, Pe��a-Santiago, Reyes & Abolafia, Joaqu��n, 2016, Re-description of Cephalobus topali Andr��ssy, 1970 (Rhabditida, Cephalobidae) from Vietnam, and transfer to Acrobeloides (Cobb, 1924) Thorne, 1937, pp. 593-600 in Zootaxa 4092 (4) on pages 594-599, DOI: 10.11646/zootaxa.4092.4.9, http://zenodo.org/record/265685

Cyrtodactylus calamei sp. nov. (Fig. 7) Holotype. VNUF R. 2015.28, adult male, in the karst forest, Tham Nok Aen region, Thong Xam Village (17 �� 34.179 ���N, 105 �� 50.329 ���E, elevation 210 m a.s.l.) within Hin Nam No NPA, Khammouane Province, central Laos, was collected on 25 March 2015 by V. Q. Luu, T. Calame, and K. Thanabuaosy. Paratypes. IEBR A. 2015.36, adult male; NUOL R- 2015.22, subadult male; VNUF R. 2015.27, adult female, the same data as the holotype. Diagnosis. Cyrtodactylus calamei sp. nov. can be distinguished from its congeners by a unique combination of the following characters: Adult SVL 80.0 �� 8.0 mm (mean �� SD); head dorsally with grey small scattered spots; nuchal loop present with indentations, not enlarged posteriorly, extending from each postnasal cross orbit and contacting on nape; four greyish brown, wide transverse bands between limbs, sometimes irregular; dorsal surface with homogenous, low, round, weakly keeled scales; 39���42 ventral scales at midbody; ventrolateral skin folds well-defined; 183���197 ventral scale rows from mental to cloacal slit; 101���114 scale rows at midbody; 35���39 precloacal-femoral pores in males, 38 in the female; enlarged femoral and precloacal scales present; four postcloacal tubercles; subcaudal scales transversely enlarged. Description of holotype. Adult male, medium sized, SVL 75.8 mm; body long (TrunkL/SVL 0.44); head moderate (HL/SVL 0.29), narrow (HW/HL 0.77), somewhat depressed (HH/HL 0.42), differentiated from neck; prefrontal and postnasal regions concave; snout elongate (SE/HL 0.45), obtuse; snout scales small, homogeneous, granular, about two times larger than those in frontal and parietal regions; eye large (OD/HL 0.29), pupils vertical; supraciliaries with tiny spines posteriorly; ear opening oval, obliquely directed, small in size (EarL/HL 0.10); rostral subrectangular, wider than high (RH/RW 0.61), medially deep furrow, vertical suture, bordered by nasorostral, nare and first supralabial laterally; nares oval, surrounded by rostral anteriorly, supranasal, first supralabial laterally, and posteriorly by two enlarged postnasals; intersupranasal scale single; mental subtriangular, nearly as wide as rostral (MW/RW 0.94), bordered by two postmentals and first infralabial laterally; supralabials nine; infralabials eight. Dorsal scales granular to flattened; dorsal tubercles round, weakly keeled, extending from postocciput to base of tail; ventrolateral folds distinct; ventral scales smooth, medial scales about two times larger than dorsal scales, imbricate, 40 rows at midbody between folds; midbody scale rows 104; ventral scales from mental to cloacal slit 183; enlarged femoral-precloacal scales present; precloacal-femoral pores 39, in a continued row; precloacal groove absent. Fore and hind limbs moderately long (ForeL/SVL 0.18, CrusL/SVL 0.2); tubercles on dorsum of fore limbs absent; dorsal surface of hind limbs interspersed with tubercles; interdigital webbing weakly developed; subdigital lamellae on fourth fingers 16 / 16 and on fourth toes 19 / 18. Tail tapering to a point (TaL/SVL 1.42); four postcloacal tubercles laterally; subcaudals distinctly enlarged. Coloration in life. Ground color of dorsal head, back, limbs, and tail yellowish brown; dorsal head with small spread spots and a heart-shaped marking on postocciput; nuchal loop present, in U-shape, from posterior corner of nare crossing eye and tympanum, extending to nape, dark brown, bolder from postocular to nape, irregularly edged in yellow; four greyish brown body bands between limb insertions with indentations in mid-dorsal region, edged in yellow; dorsal surface of fore and hind limbs with grey reticulations; tail with narrow light bands; ventral surface greyish cream. Sexual dimorphism. The single adult female differs from two adult males by its larger size (maximum SVL 89.3 mm versus 75.8 mm in the males) and lacking of hemipenial swellings at the base of tail (see Table 4 & Fig. 6). Comparisons. We compared Cyrtodactylus calamei sp. nov. with other Cyrtodactylus species known from Laos and neighbouring countries in the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on examination of specimens (see Appendix) and data provided from taxonomic publications (Luu et al. 2014 a; Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014; Sumontha et al. 2015; Nguyen et al. 2015; Luu et al. 2015 a; Luu et al. 2016 a, b) (see Table 3). The cluster and correspondence analyses indicated that Cyrtodactylus calamei sp. nov. is nested in the same clade with C. darevskii and the species to be described in the following (Figs. 2���3). Molecular phylogenetic analyses also strongly supported the sister relationship between the new species and afore mentioned taxa (see Fig. 1). Morphologically, Cyrtodactylus calamei sp. nov. closely resembles the other karst forest species, C. darevskii and C. phongnhakebangensis, in dorsal colour pattern. However, the new species can be distinguished from C. darevskii by its smaller size (maximum SVL 89.3 mm versus 100 mm), having fewer dorsal tubercle rows (10���16 versus 16���20), fewer femoral and precloacal pores in males (35���39 versus 38���44), more femoral and precloacal pores in females (38 versus 24���34), the presence of heart-shaped marking on postocciput (versus absent), four greyish brown regular transverse body bands as wide as nearly two times of nuchal band (versus four to five dark irregular transverse breaking bands as narrow as nuchal band), first body band wide, butterfly-shaped (versus thin, U-shaped in C. darevskii), tail with light rings (versus banded); from C. phongnhakebangensis by its smaller size (maximum SVL 89.3 mm versus 96.3 mm), more scale rows from mental to the front of cloacal slit (183���197 versus 161���177), nuchal loop narrow, indented, not posteriorly enlarged (versus wide, posteriorly enlarged), four greyish brown transverse body bands, slightly narrower light bands (versus three to five dark transverse body bands, twice wider than light bands), and tail with light rings (versus bands) (see Table 4). Distribution. Cyrtodactylus calamei is currently known only from the type locality in Tham Nok En area, Hin Nam No NPA, Khammouane Province, central Laos (Fig. 6). Etymology. The new species is named in honour of our friend and colleague, Mr. Thomas Calame, from WWF Greater Mekong, Vientiane, Laos, who participated in our field research in Hin Nam No NPA, Khammouane Province between 2014 and 2015. As common names, we suggest Calame���s Bent-toed Gecko (English), Ki Chiem Calame (Laotian). Natural history. Specimens were found at night between 19: 30 and 21:08h, on limestone outcrops, at elevations between 190 and 260 m a.s.l. The surrounding habitat was karst forest. The relative humidity was 80 % and the air temperature ranged from 23 to 26 o C (see Table 5)., Published as part of Luu, Vinh Quang, Bonkowski, Michael, Nguyen, Truong Quang, Le, Minh Duc, Schneider, Nicole, Ngo, Hanh Thi & Ziegler, Thomas, 2016, Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos, pp. 101-140 in Zootaxa 4107 (2) on pages 113-117, DOI: 10.11646/zootaxa.4107.2.1, http://zenodo.org/record/266526, {"references":["Luu, V. Q., Calame, T., Bonkowski, M., Nguyen, T. Q. & Ziegler, T. (2014 a) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Khammouane Province, Laos. Zootaxa, 3760 (1), 54 - 66. http: // dx. doi. org / 10.11646 / zootaxa. 3760.1.3","Nazarov, R. 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. http: // dx. doi. org / 10.11646 / zootaxa. 3784.1.2","Schneider, N., Nguyen, T. Q., Le, M. D., Nophaseud, L., Bonkowski, M. & Ziegler, T. (2014) A new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of northern Laos. Zootaxa, 3835 (1), 080 - 096. http: // dx. doi. org / 10.11646 / zootaxa. 3835.1.4","Sumontha, M., Pauwels, O. S. G., Panitvong, N., Kunya, K. & Grismer, J. L. (2015) A new lowland forest Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Ranong Province, peninsular Thailand. Zootaxa, 3911 (1), 106 - 118. http: // dx. doi. org / 10.11646 / zootaxa. 3911.1.6","Le, M., Nguyen, H. M., Duong, H. T., Nguyen, T. V., Dinh, L. D., Nguyen, N. X., Nguyen, L. D., Dinh, T. H. & Nguyen, D. X. (2015) Phylogeography of the Laotian rock rat (Diatomyidae: Laonastes): implications for Lazarus taxa. Mammal Study, 40, 109 - 114. http: // dx. doi. org / 10.3106 / 041.040.0206","Luu, V. Q., Calame, T., Nguyen, T. Q., Bonkowski, M. & Ziegler, T. (2015 a) A new species of Cyrtodactylus (Squamata: Gekkonidae) from limestone forest, Khammouane Province, central Laos. Zootaxa, 4058 (3), 388 - 402. http: // dx. doi. org / 10.11646 / zootaxa. 4058.3.6","Luu, V. Q., Calame, T., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016 a) Cyrtodactylus rufford, a new cavedwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4067 (2), 185 - 199. http: // doi. org / 10.11646 / zootaxa. 4067.2.4","Luu, V. Q., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016 b) A new species of karst-dwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4079 (1), 087 - 102. http: // doi. org / 10.11646 / zootaxa. 4079.1.6"]}

Cyrtodactylus wayakonei group Species. C. spelaeus, C. vilaphongi, C.wayakonei. Characters. Adult SVL 86.1 ���91.0 mm, supranasals 0���1; dorsal tubercles 10���19; webbing between fingers and toes absent; tubercles on limbs present; a low number of precoacal pores in males 6���8; postcloacal tubercles 2���4; subcaudals slightly enlarged; dorsal body blotched (except for C. spelaeus). Distribution. The group is distributed in karst formations of northern Laos including Luang Nam Tha, Luang Prabang, and Vientiane provinces., Published as part of Luu, Vinh Quang, Bonkowski, Michael, Nguyen, Truong Quang, Le, Minh Duc, Schneider, Nicole, Ngo, Hanh Thi & Ziegler, Thomas, 2016, Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos, pp. 101-140 in Zootaxa 4107 (2) on page 135, DOI: 10.11646/zootaxa.4107.2.1, http://zenodo.org/record/266526

Cyrtodactylus species groups in Laos As currently recognized, 21 Cyrtodactylus species are known from Laos. Based on our molecular phylogeny, the molecular data presented in Nazarov et al. (2014), and a combination of characters including size, dorsal color pattern, scalation, and geographic distribution, we herein propose four species groups within the genus Cyrtodactylus occurring in Laos (see Table 13), which are defined as follows, Published as part of Luu, Vinh Quang, Bonkowski, Michael, Nguyen, Truong Quang, Le, Minh Duc, Schneider, Nicole, Ngo, Hanh Thi & Ziegler, Thomas, 2016, Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos, pp. 101-140 in Zootaxa 4107 (2) on page 132, DOI: 10.11646/zootaxa.4107.2.1, http://zenodo.org/record/266526, {"references":["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."]}

Cyrtodactylus sommerladi sp. nov. (Fig. 10) Holotype. VNUF R. 2013.22 adult male, in karst forest, Hang Toi region, Noong Ma Village (17 �� 17.795 ���N, 106 ��08.738���E, elevation 572 m a.s.l.) within Hin Nam No NPA, Khammouane Province, central Laos, collected on 0 5 May 2013 by V. Q. Luu and N. V. Ha. Paratypes. NUOL R- 2013.23, IEBR A. 2015.37, ZFMK 97196, VNUF R. 2013.105, VNUF R. 2014.87, NUOL R- 2013.14, IEBR A. 2015.39, NUOL R- 2013.21, IEBR A. 2015.38, VNUF R. 2013.104, ZFMK 97197, VNUF R. 2014.89, VNUF R. 2013.67 the same locality as the holotype. IEBR A. 2015.40 adult female, in karst forest, Cha Lou Village (17 �� 18.880 ���N, 105 �� 57.103 ���E, elevation 572 m a.s.l.) within Hin Nam No NPA, Khammouane Province, central Laos, collected on 25 May 2013. Diagnosis. Cyrtodactylus sommerladi sp. nov. is characterized by: Adult SVL 72.3 �� 3.8 mm (mean �� SD); dorsal head greyish brown without dark blotches; nuchal loop present, narrow, not enlarged posteriorly; five or six dark transverse bands between limbs; dorsal surface with homogenous, tubercle-like scales; ventral scales at midbody 31���39; ventrolateral skin folds present; ventral scale rows from mental to cloacal slit 168���192; scale rows at midbody 76���93; precloacal-femoral pores 20���26 in males, 17���21 in females; enlarged femoral and precloacal scales present; postcloacal tubercles 4���6; subcaudal scales slightly enlarged. Description of holotype. Adult male, small sized (SVL 70.3 mm); body elongate (TrunkL/SVL 0.45); head elongate (HL/SVL 0.25), width (HW/HL 0.77), relatively depressed (HH/HL ratio 0.41), distinct from neck; loreal area concave; snout long (SE/HL ratio 0.47), obtuse, two times longer than diameter of orbit (OD/SE 0.50); snout scales small, homogeneous, granular, about one and a half times larger than those in frontal and parietal regions; eye large (OD/HL ratio 0.23), pupils vertical; eyelid fringe with tiny spines in posterior part, posterior ones more developed; ear opening oval, obliquely orientated, small (EarL/HL 0.12); rostral square-shaped, wider than high (RH/RW ratio 0.55), medially with a straight, vertical suture, in contact with nasorostral, nare and first supralabial on each side; nares oval, surrounded by rostral anteriorly, supranasal, first supralabial laterally, and two enlarged postnasals posteriorly; supranasals in contact; mental triangular, slightly narrower than rostral (RW/MW ratio 0.94), in contact with two postmentals and the first infralabial on each side, postmentals surrounded by first infralabial on each side and seven granular scales posteriorly, two outer ones enlarged; supralabials 10 / 10; infralabials 8 / 9; supralabials separated from orbit by 3 or 4 rows of granular scales. Dorsal scales homogenous, tubercle-like; dorsal tubercles indistinct; ventrolateral folds present; ventral scales smooth, medial scales 2 or 3 times larger than dorsal scales, round, and in 37 rows at midbody; midbody scale rows 87; scales between mental and cloacal slit 192; femoral scales enlarged; precloacal scales enlarged, precloacal-femoral pores 23, in a continuous row; precloacal groove absent. Fore and hind limbs moderately slender (ForeL/SVL 0.15, CrusL/SVL 0.19); dorsal surface of hind limbs with tubercles, absent on fore limbs; fingers and toes with rudimental webbing; lamellae under fourth fingers 18 / 18, under fourth toes 20 / 22. Tail longer than snout-vent length (TaiL/SVL 1.14); postcloacal tubercles 6 on each side; subcaudals slightly enlarged. ......continued on the next page Coloration in life. Ground color of dorsal head and back greyish brown; head without dark blotches; nuchal loop present, narrow, in U-shape, not enlarged posteriorly, extending from posterior corner of eye through tympanum to the neck, dark brown, edged in yellow posteriorly; four dark brown body bands between limb insertions, edged in yellow; dorsal surface of fore and hind limbs with greyish-brown dark spots; tail dorsally brown with 12 yellow-whitish rings, edged in yellow; ventral surface bright beige. Variation. The nuchal loop in the adult female paratype (ZFMK 97197) is interrupted in the middle. Other paratypes have some indistinct spot markings on dorsal head and somewhat irregular and breaking transverse bands. Sexual dimorphism. The females differ from the males by having fewer precloacal-femoral pores (17���21 versus 20���26 in the males) and females lack hemipenial swellings at the tail base (see Table 9). Comparisons. We compared the new species with its congeners from Laos and neighbouring countries in the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on data obtained from the literature (compiled after Luu et al. 2014 a; Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014; Sumontha et al. 2015; Nguyen et al. 2015; Luu et al. 2015 a; Luu et al. 2016 a, b) and based on the examination of specimens and photographs (see Appendix and Table 3). Cyrtodactylus sommerladi sp. nov. was well separated by the cluster and correspondence analyses from C. roesleri (Figs. 2���3). Molecular phylogenetic analyses also revealed the close relationships between these species (see Fig. 1). The new species is most similar to C. roesleri from Vietnam in body size, and dorsal pattern. However, the new species can be distinguished from the latter by its larger size (maximal SVL 80.3 mm versus 73.5 mm), having more dark body bands: 5 or 6 (n= 15) versus 4���5 (n= 19), dorsal surface of the new species shows homogenous scales, all appear tubercle-like, with unclear dorsal tubercles (0���5) (versus dorsal tubercles being arranged in 13���19 rows) (Fig. 11), dorsal surface of tail without tubercles (versus present) (see Fig. 12 A,B) and the shape of the rostral suture in the new species is just straight vertical versus Y-shaped in C. roesleri. In addition, Cyrtodactylus sommerladi sp. nov. can be distinguished from C. roesleri by the different head shape, the head of the new species is wider and flatter (Fig. 5). Distribution. Currently, Cyrtodactylus sommerladi sp. nov. is known only from the type locality in the karst forest of Hin Nam No NPA, Khammouane Province, central Laos (Fig. 6). Etymology. The specific epithet sommerladi refers to our colleague and good friend Ralf Sommerlad, late Regional vice chairman of the IUCN SSC Crocodile Specialist Group (CSG) for Europe, who passed away on 11 June 2015, to honor his lifework and strong commitment for reptile conservation. As common names, we suggest Sommerlad���s Bent-toed Gecko (English), Ki Chiem Sommerlad (Laotian). Natural history. Specimens were collected at night from 19: 20 to 21: 43 h, mainly on karst walls, ca. 1���4.5 m above the ground, near cave entrances in the limestone forest, at elevations between 269 and 614 m a.s.l. Two female specimens IEBR A. 2015.38 and ZFMK 97197 were found on tree trunks, about 1���2 m from the forest floor. The surrounding habitat was karst forest dominated by species of Ebenaceae, Dracaenaceae, Arecaeae, Poaceae, Meliaceae, and Moraceae. The relative humidity was between 74 % and 90 %, and temperatures ranged from 22.7 to 29.9 o C (see Table 10). Female specimens contained eggs in May, by contrast, no records of gravid females or hatchlings were made in March. Thus, the breeding season seems to start only in April or May. The majority of the caves, karst walls, and karst forests where Cyrtodactylus sommerladi sp. nov. have been explored had dry surfaces, without flowing streams inside (e.g., Ellis & Pauwels 2012)., Published as part of Luu, Vinh Quang, Bonkowski, Michael, Nguyen, Truong Quang, Le, Minh Duc, Schneider, Nicole, Ngo, Hanh Thi & Ziegler, Thomas, 2016, Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos, pp. 101-140 in Zootaxa 4107 (2) on pages 123-128, DOI: 10.11646/zootaxa.4107.2.1, http://zenodo.org/record/266526, {"references":["Luu, V. Q., Calame, T., Bonkowski, M., Nguyen, T. Q. & Ziegler, T. (2014 a) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Khammouane Province, Laos. Zootaxa, 3760 (1), 54 - 66. http: // dx. doi. org / 10.11646 / zootaxa. 3760.1.3","Nazarov, R. 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. http: // dx. doi. org / 10.11646 / zootaxa. 3784.1.2","Schneider, N., Nguyen, T. Q., Le, M. D., Nophaseud, L., Bonkowski, M. & Ziegler, T. (2014) A new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of northern Laos. Zootaxa, 3835 (1), 080 - 096. http: // dx. doi. org / 10.11646 / zootaxa. 3835.1.4","Sumontha, M., Pauwels, O. S. G., Panitvong, N., Kunya, K. & Grismer, J. L. (2015) A new lowland forest Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Ranong Province, peninsular Thailand. Zootaxa, 3911 (1), 106 - 118. http: // dx. doi. org / 10.11646 / zootaxa. 3911.1.6","Le, M., Nguyen, H. M., Duong, H. T., Nguyen, T. V., Dinh, L. D., Nguyen, N. X., Nguyen, L. D., Dinh, T. H. & Nguyen, D. X. (2015) Phylogeography of the Laotian rock rat (Diatomyidae: Laonastes): implications for Lazarus taxa. Mammal Study, 40, 109 - 114. http: // dx. doi. org / 10.3106 / 041.040.0206","Luu, V. Q., Calame, T., Nguyen, T. Q., Bonkowski, M. & Ziegler, T. (2015 a) A new species of Cyrtodactylus (Squamata: Gekkonidae) from limestone forest, Khammouane Province, central Laos. Zootaxa, 4058 (3), 388 - 402. http: // dx. doi. org / 10.11646 / zootaxa. 4058.3.6","Luu, V. Q., Calame, T., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016 a) Cyrtodactylus rufford, a new cavedwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4067 (2), 185 - 199. http: // doi. org / 10.11646 / zootaxa. 4067.2.4","Luu, V. Q., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016 b) A new species of karst-dwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4079 (1), 087 - 102. http: // doi. org / 10.11646 / zootaxa. 4079.1.6","Ellis, M. & Pauwels, O. S. G. (2012) The bent-toed geckos (Cyrtodactylus) of the caves and karst of Thailand. Cave and Karst Science, 39 (1), 16 - 22."]}

Cyrtodactylus hinnamnoensis sp. nov. (Fig. 8) Cyrtodactylus phongnhakebangensis��� Luu, Nguyen, Calame, Hoang, Soudthichack, Bonkowski, Ziegler, 2013. Biodiversity Data Journal, e 1015: 4. Holotype. IEBR A. 2013.90, adult male, from Ban Dou Village (17 �� 30.385 ���N, 105 �� 49.160 ���E, elevation 183 m a.s.l.) within Hin Nam No NPA, Khammouane Province, central Laos, was collected on 11 June 2013 by V. Q. Luu and N. V. Ha. Paratypes. IEBR A. 2013.89, adult male, 7 May 2013, from Hang Toi region, Noong Ma Village (17 �� 17.766 ���N, 106 ��08.803���E, elevation 580 m a.s.l.); VNUF R. 2013.1 and NUOL R- 2013.2, adult males, 9 June 2013, from Vang Ma No Village (17 �� 30.778 ���N, 105 �� 49.259 ���E, elevation 180 m a.s.l.); VNUF R. 2014.99, adult male, 27 May 2014, from Cha Lou Village (17 �� 19.504 ���N, 105 �� 57.630 ���E, elevation ca. 300 m a.s.l.); ZFMK 95235, adult female, 8 May 2013, from Hang Toi region, Noong Ma Village (17 �� 17.763 ���N, 106 ��08.778���E, elevation 555 m a.s.l.); ZFMK 95236, adult female, 30 May 2013, from Noong Choong Region, Cha Lou Village (17 �� 20.248 ���N, 105 �� 56.693 ���E, elevation 252 m a.s.l.); NUOL R- 2013.3, adult female, 11 June 2013, from Ban Dou Village (17 �� 31.545 ���N, 105 �� 49.086 ���E, elevation 197 m a.s.l.); VNUF R. 2015.3, female, 13 March 2015, from Xebangfai cave, Noong Ping Village (17 �� 22.459 ���N, 105 �� 49.626 ���E, elevation 182 m a.s.l.); NUOL R- 2015.9, female, 13 March 2015, from Xebangfai cave, Noong Ping Village (17 �� 22.648 ���N, 105 �� 52.931 ���E, elevation 182 m a.s.l.); VNUF R. 2015.11, female, 14 March 2015, from Xebangfai cave, Noong Ping Village (17 �� 22.759 ���N, 105 �� 52.931 ���E, elevation 285 m a.s.l.). The paratypes (VNUF R. 2015.3, NUOL R- 2015.9, and VNUF R. 2015.11) were collected by V. Q. Luu and K. Thanabuaosy in March 2015; the paratype (VNUF R. 2014.99) was collected by V. Q. Luu, N. V. Ha, T. Calame, D. V. Phan and K. Thanabuaosy in May 2014, the remaining type series was collected by V. Q. Luu, N. V. Ha, and K. Thanabuaosy in May and June 2013 (V. Q. Luu et al.). Diagnosis. Cyrtodactylus hinnamnoensis sp. nov. is characterized by: Adult SVL 84.1 �� 11.7 mm (mean �� SD); dorsal head with dark blotches; nuchal loop wide, distinct, posteriorly enlarged; dorsal body with four to six blackish brown bands between limb insertions; 13���19 irregular, weakly keeled dorsal tubercle rows; 35���48 ventral scale rows; ventral scale rows from mental to cloacal slit 179���201; scale rows at midbody 93���112; ventrolateral folds present, without tubercles; 36���44 precloacal-femoral pores in the males; 0���28 pores in females; enlarged femoral and precloacal scales present; 4���6 postcloacal tubercles; subcaudals enlarged. Description of holotype. Adult male, snout-vent length (SVL) 83.6 mm; body elongate (TrunkL/SVL 0.42); head elongate (HL/SVL 0.27), relatively wide (HW/HL 0.68), depressed (HH/HL 0.40), distinguished from neck; loreal region concave; snout long (SE/HL 0.43), obtuse, longer than diameter of orbit (OD/SE 0.60); snout scales small, homogeneous, granular, larger than those on frontal and parietal regions; eye large (OD/HL 0.26), pupils vertical; eyelid fringe with tiny spines posteriorly; ear oval-shaped, small (EarL/HL 0.08); rostral wider than high (RH/RW 0.60), with a median suture; supranasals in contact anteriorly and separated from each other by a small scale posteriorly; rostral in contact with first supralabial and nostril scales on each side; nares oval, surrounded by supranasal, rostral, first supralabial, and two enlarged postnasals; mental triangular, wider than high (ML/MW 0.65); two enlarged postmentals in broad contact posteriorly, bordered by mental anteriorly, first two infralabials laterally, and eight small scales posteriorly; supralabials 12 / 10; infralabials 10 / 9. Dorsal scales granular to flattened; dorsal tubercles round, weakly keeled, present on occiput, back and tail base, each surrounded by 8 granular scales, in 15 irregular longitudinal rows at midbody; ventral scales smooth, medial scales 2���3 times larger than dorsal scales, round, subimbricate, largest posteriorly, in 35 longitudinal rows between lateral folds at midbody; ventrolateral folds present, without tubercles; gular region with homogeneous smooth scales; ventral scales in a line from mental to cloacal slit 186; precloacal groove absent; enlarged femoral scales present; femoral and precloacal pores 42. Fore and hind limbs moderately slender (ForeL/SVL 0.17, CrusL/SVL 0.21); dorsal fore limbs with slightly developed tubercles; dorsal hind limbs covered by distinctly developed tubercles; fingers and toes free of webbing; lamellae under fourth fingers 19 / 19, under fourth toes 20 / 20. Tail regenerated, postcloacal tubercles 5 / 5; dorsal tail bearing tubercles at base; subcaudals distinctly enlarged, flat, smooth. Coloration in life. Ground coloration of dorsal head greyish brown with dark blotches; nuchal loop black, in U-shape, from posterior corner of eye through tympanum to the neck, dark brown, edged in yellow; body bands between limb insertions five, somewhat irregular, dark brown, edged in white; dorsal surface of fore and hind limbs with dark bars; tail brown dorsally with light brown rings, edged by yellowish white; chin, throat, and belly cream; upper and lower lips with dark brown bars; tail ventrally grey with light dots. Sexual dimorphism. The females differ from the males by lacking or having fewer precloacal-femoral pores (0���28 versus 36���44 in the males) and the absence of hemipenial swellings at the tail base (see Table 6). Comparisons. We compared the new species with its congeners from Laos and neighbouring countries in the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on examination of specimens (see Appendix) and data integrated from the literature (compiled after Luu et al. 2014 a; Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014; Sumontha et al. 2015; Nguyen et al. 2015; Luu et al. 2015 a; Luu et al. 2016 a, b) (see Tables 3). The cluster and correspondence analyses of morphological characters supported Cyrtodactylus hinnamnoensis sp. nov. as a sister taxon to C. darevskii (Figs. 2���3). Molecular phylogenetic analyses also demonstrated the close relationships between these species (see Fig. 1). Morphologically, Cyrtodactylus hinnamnoensis sp. nov. is closely related to the C. phongnhakebangensis group including C. darevskii, C. phongnhakebangensis, C. calamei by dorsal colour pattern and the number of cloacal and femoral pores in males. However, the new species can be distinguished from C. darevskii by having fewer cloacal and femoral pores in females (maximum 0���28 versus 24���34), four to six blackish brown transverse body bands, as wide as light bands (versus four to five dark irregular transverse breaking body bands, 0.5 times narrower than light band), first body band wide, butterfly-shaped (versus thin, U-shaped in C. darevskii), the presence of tubercles on fore limbs (versus absent), and tail consisting of light rings (versus banded); Cyrtodactylus hinnamnoensis sp. nov. differs from C. phongnhakebangensis by its slightly larger size (SVL reaching 100.6 mm versus 96.3 mm), having fewer cloacal and femoral pores in females (0���28 versus 0���41), having more scale rows from mental to the front of cloacal slit (179���201 versus 161���177), the presence of tubercles on fore limbs (versus absent), a narrower nuchal loop, not enlarged posteriorly (wide, enlarged posteriorly), four to six blackish brown transverse body bands as wide as light bands (versus three to five dark transverse body bands as wide as double light bands, light transverse bands with small spots (versus with big black blotches), and tail pattern consisting of light rings (versus banded); Cyrtodactylus hinnamnoensis sp. nov. differs from C. calamei by its larger size (SVL reaching 100.6 mm versus 89.3 mm), fewer cloacal and femoral pores in females (0���28 versus 38), more postcloacal tubercles (4���6 versus 4), dorsal head marking with distinctly dark spots and blotches (versus indistinct dots), the absence of heart-shaped marking on postocciput (versus present), four to six blackish brown body = standard deviati��n, f��r ��ther abbreviati��ns see material and meth��ds). ������continued on the next page transverse bands, as wide as light bands (versus four greyish brown transverse bands, narrower than light bands) (for more details see Table 7, Fig. 9). The results of the correspondence analysis comparing all adult male morphological maesurements of Cyrtodactylus hinnamnoensis sp. nov. and the latter species indicated four distinct groups between these species (see Fig. 4). Distribution. Cyrtodactylus hinnamnoensis sp. nov. is currently known only from the type locality in the karst forest of Hin Nam No NPA, Khammouane Province, central Laos (Fig. 6). Etymology. We name this species after its type locality, Hin Nam No NPA where the new Cyrtodactylus species was discovered and propose the following common names: Hinnamno Bent-toed Gecko (English), Ki Chiem Hin Nam No (Laotian). Natural history. Specimens were found at night between 19: 41 and 22:03h on karst walls, ca. 0.3���5 m above the ground, near cave entrances in the limestone forest, at elevations between 175 and 580 m a.s.l. Only one male specimen VNUF R. 2014.99 was collected on a tree trunk, about 1 m from the forest floor. The surrounding habitat was karst forest, dominated by species of Ebenaceae, Dracaenaceae, Arecaeae, Poaceae, Meliaceae, and Moraceae. The relative humidity ranged from 78 % to 90 %, and temperatures were from 24.9 to 30.7 o C (see Table 8). When capturing individuals of the species, we observed an increased rate of tail autotomy and many individuals had regenerated tail, for example, seven of 11 specimens of Cyrtodactylus hinnamnoensis sp. nov. had dropped or/and regenerated tails. This suggests that these populations might be under the stress of predators (see also Grismer et al. 2016)., Published as part of Luu, Vinh Quang, Bonkowski, Michael, Nguyen, Truong Quang, Le, Minh Duc, Schneider, Nicole, Ngo, Hanh Thi & Ziegler, Thomas, 2016, Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos, pp. 101-140 in Zootaxa 4107 (2) on pages 117-123, DOI: 10.11646/zootaxa.4107.2.1, http://zenodo.org/record/266526, {"references":["Luu, V. Q., Nguyen, T. Q., Calame, T., Hoang, T. T., Southichack, S., Bonkowski, M. & Ziegler, T. (2013) New country records of reptiles from Laos. Biodiversity Data Journal, 1, 1 - 14. http: // dx. doi. org / 10.3897 / BDJ. 1. e 1015","Luu, V. Q., Calame, T., Bonkowski, M., Nguyen, T. Q. & Ziegler, T. (2014 a) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Khammouane Province, Laos. Zootaxa, 3760 (1), 54 - 66. http: // dx. doi. org / 10.11646 / zootaxa. 3760.1.3","Nazarov, R. 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. http: // dx. doi. org / 10.11646 / zootaxa. 3784.1.2","Schneider, N., Nguyen, T. Q., Le, M. D., Nophaseud, L., Bonkowski, M. & Ziegler, T. (2014) A new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of northern Laos. Zootaxa, 3835 (1), 080 - 096. http: // dx. doi. org / 10.11646 / zootaxa. 3835.1.4","Sumontha, M., Pauwels, O. S. G., Panitvong, N., Kunya, K. & Grismer, J. L. (2015) A new lowland forest Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Ranong Province, peninsular Thailand. Zootaxa, 3911 (1), 106 - 118. http: // dx. doi. org / 10.11646 / zootaxa. 3911.1.6","Le, M., Nguyen, H. M., Duong, H. T., Nguyen, T. V., Dinh, L. D., Nguyen, N. X., Nguyen, L. D., Dinh, T. H. & Nguyen, D. X. (2015) Phylogeography of the Laotian rock rat (Diatomyidae: Laonastes): implications for Lazarus taxa. Mammal Study, 40, 109 - 114. http: // dx. doi. org / 10.3106 / 041.040.0206","Luu, V. Q., Calame, T., Nguyen, T. Q., Bonkowski, M. & Ziegler, T. (2015 a) A new species of Cyrtodactylus (Squamata: Gekkonidae) from limestone forest, Khammouane Province, central Laos. Zootaxa, 4058 (3), 388 - 402. http: // dx. doi. org / 10.11646 / zootaxa. 4058.3.6","Luu, V. Q., Calame, T., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016 a) Cyrtodactylus rufford, a new cavedwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4067 (2), 185 - 199. http: // doi. org / 10.11646 / zootaxa. 4067.2.4","Luu, V. Q., Nguyen, T. Q., Le, M. D., Bonkowski, M. & Ziegler, T. (2016 b) A new species of karst-dwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4079 (1), 087 - 102. http: // doi. org / 10.11646 / zootaxa. 4079.1.6","Grismer, L. L., Wood, P. L. Jr., Anuar, S., Davis, H. R., Burch, B. T., Cobos, A. J. & Murdoch, M. L. (2016) A new species of karst forest Bent-toed Gecko (genus Cyrtodactylus Gray) not yet threatened by foreign cement companies and a summary of Peninsular Malaysia'sendemic karst forest herpetofauna and the need for its conservation, Zootaxa, 4061 (1), 1 - 17. http: // doi. org / 10.11646 / zootaxa. 4061.1.1"]}

Nguyen, Thi Anh Duong, Abolafia, Joaquín, Bonkowski, Michael, Santiago, Reyes Peña- (2016): Two new species of the genus Aporcelinus Andrássy, 2009 (Nematoda, Dorylaimida, Aporcelaimidae) from Vietnam. Zootaxa 4103 (6): 550-560, DOI: http://doi.org/10.11646/zootaxa.4103.6.5

Luu, Vinh Quang, Bonkowski, Michael, Nguyen, Truong Quang, Le, Minh Duc, Schneider, Nicole, Ngo, Hanh Thi, Ziegler, Thomas (2016): Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos. Zootaxa 4107 (2): 101-140, DOI: http://doi.org/10.11646/zootaxa.4107.2.1

Odorrana mutschmanni sp. nov. (Figs. 2, 3) Holotype: IEBR 3723 (Field No. CB 2015.12), adult male, collected by T.Q. Nguyen on 22 April 2015 in the karst forest near Coong Village (22o42.712’N, 106o40.075’E, at an elevation of 447 m), Duc Quang Commune, Ha Lang District, Cao Bang Province, Vietnam. Paratypes: IEBR 3724 (Field No. CB 2012.77), adult males, collected on 15 April 2012; IEBR 3725 (Field No. CB 2012.89), adult female and IEBR 3726–3729 (Field No. CB 2012.90–93), adult males, collected on 16 April 2012, by T.Q. Nguyen et al.; ZFMK 97329, 97330 (Field No. CB 2012.139, 2012.140), adult males, collected on 3 May 2012, by H. T. An, S. Herbst and T. Lehmann; IEBR 3730 (Field No. CB2014.16), adult female, collected by C.T. Pham et al. on 10 June 2014, IEBR 3731 (Field No. CB 2015.11), adult female, collected by T.Q. Nguyen on 22 April 2015, the same data as the holotype. Diagnosis. The new species was strongly supported as a member of Odorrana based on molecular analyses (Fig. 1) and is distinguishable from its congeners by a combination of the following morphological characters: (1) size large (SVL 85.9–91.6 mm in males, 108.7–110.1 mm in females); (2) head longer than wide; (3) vomerine teeth present; (4) external vocal sacs absent; (5) snout short (SL/SVL 0.16–0.17); (6) tympanum large (TD/ED 0.70 in males, 0.68 in females); (7) dorsal surface of head and anterior part of body smooth, posterior part of body and flanks with small tubercles; (8) supratympanic fold present; (9) dorsolateral fold absent; (10) webbing formula I0– 0II0–0III0–1/ 2IV 1/2–0V; (11) in life, dorsum green with dark brown spots; (12) flanks greyish brown with dark brown spots; (13) throat and chest grey, underside of limbs with large dark brown spots, edged in white, forming a network. Description of holotype. Adult male; SVL 85.9 mm; head longer than wide (HL 33.6 mm, HW 29.9 mm); snout round anteriorly in dorsal view, projecting beyond lower jaw; nostril lateral, closer to the snout tip than to eye (NS 6.2 mm, EN 7.5 mm); canthus rostralis distinct; pupil horizontally oval; loreal region slightly concave and oblique; snout length greater than eye diameter (SL 14.0 mm, ED 9.9 mm); internarial distance wider than interorbital distance and upper eyelid (IND 10.5. mm, IOD 9.7 mm, UEW 6.6 mm); tympanum distinct, round, 70% eye diameter (TD 6.9 mm); vomerine teeth in two oblique ridges; tongue cordiform, deeply notched posteriorly; vocal sac absent. Forelimbs: Forelimb length (FLL 16.6 mm), hand length (HAL 44.4 mm); relative finger lengths: II 2 times of the width of phalanges and about 40% the diameter of tympanum; subarticular tubercles round, formula 1, 1, 2, 2; inner metatarsal tubercle oval, elongate; outer metatarsal tubercle small; finger I with nuptial pad, elongate. Hindlimbs: Tibia longer than thigh (FeL 38.1 mm, TbL 46.9 mm), approximately five times longer than wide (TbW 9.8 mm); tips of toes expanded into discs, with circummarginal grooves; width of toe IV disc narrower than width of finger III disc, approximately two times of the width of phalanges; relative length of toes: I Skin: Dorsal surface of head and anterior part of body smooth; posterior part of body and flanks with tubercles; spinules present on lateral sides of body, anterior and posterior edge of tympanum; supratympanic fold present; dorsolateral fold absent; dorsal surface of limbs granular; throat, chest, belly and ventral surface of thigh smooth. Coloration in life: Iris black; dorsum green with dark brown spots; lateral side of head and flanks greyish brown with dark brown spots; lips with dark bars; tympanum dark brown; spinules on flank ivory; dorsal surface of fore and hindlimbs greyish brown with dark crossbars; throat and chest grey; ventral surface of fore and hindlimbs and belly with large dark brown spots, edged in white, forming a network; toe webbing dark brown. Coloration in alcohol: Dorsum brown, flank cream with large spots; lips with black bars; fore and hindlimbs brown with dark crossbars; throat and chest grey; belly and ventral surface of fore and hindlimbs with large black spots; toe webbing brown. Variation. Measurements and morphological characters of the type series are given in Table 3. Sexual dimorphism. The males are smaller than the females in size (SVL: 85.9–91.6 mm, n = 8 vs 108.7– 110.1 mm, n = 3, respectively). Etymology. We name this new species in honor of Dr. Frank Mutschmann, director of EXOMED in Berlin, in recognition of his support of our research and conservation work in Vietnam. As common names we suggest Mutschmann’s Frog (English), Ếch đá mut-x-man (Vietnamese), and Mutschmanns Frosch (German). Ecological notes. Odorrana mutschmanni sp. nov. appears closely associated with karst environments. Specimens were found at night between 19:00 and 23:00 h around a water pool near Coong Village, Duc Quang Commune (Fig. 4). The surrounding habitat was a secondary karst forest of medium and small hardwoods mixed Odorrana mutschmanni !" / & ’(’ () ’ * (’ (’ + ( ’ ( ’ (& ’ && ’ (0 ’ & / & & & & & & & & &’ ’ & .. % % % % % % % % % #! $ ! # $ ! # $ # $ !! $ # #! $ ­,,­ $! # # $! # $! 1 2 2 2 2 2 2 2 2 5 1 6 " 3 3 3 5 1 6 " + + & & 78 +0 * +0 + + / / +* /) * +0 +9 * ++ 6() /+ ’ / * / /+ ’9 / / 06/ ’ 8 && * &* 0 && 0 &) / &) / && &) &* & && 09&* 0 &) 0 6 () () ’)& 0) (9)& 0)(6 (: (& ’ ((/ (+ (* &/ ’ &/ * (+ 9& ’ &/ /6/ &+ ’ & 0) &+ ’9) & +6 (……continued on the next page @ !" / & () (* ’ ’ ((+ (& (&& (0 /& ( ’ & & ’ ’ & & ’ & ’ & ’ ’ & ’ & ’ & & ’.. % % % % % % % % % # $ $ # # $ $ # $ # $ # ­ ­ $ # # $ # $ !!!!!!,,!!! 1 2 2 2 2 2 2 2 2 5 1 6 " 3 3 3 5 1 6 " + + & & 8 + 0 ’ ’) ’) / ’ & ’ ’ + ’ 9 / ’ +6/ ’ ((((& 0 ((0 (((9(& 0 ((’6/ ’ 8) ’ + ’ (’ 0 + * +) * ’ * ’ (9 * + &6/ 4 * + *) 0 ’ * + * ’ * ’) * 0 0 ’9’) * *6/ 0 ’ + + (+ ’ +9+ (+ /6/ (& (’ (0 (0 (0 (’ & (((+ (09& ((’6/ (& /) * &) & /9) * & ’6/ + 8 &’ /)(& &+ ()(+ &+ &’) &’ * &+ &’ /9)(+ &+ 6(&)) 0)+ 0 0& &)) 090& &)+ +6)) 8)’ +)* +))* +)*)+)) ()*)* +9))’ +6 (0’ ’ 0 0 0 */ / 0 0 09*/ / 0’ ’6(&: 0 / + / + ’ / (/) + 0 9 / + / 6/ 0 + & ((& +9 & (()6/ ’ 8 * / *’ (** + ’/ & *) + *0 ’ *’ *0 / *) +9’/ & *’ 6 +& +&) +’ +&)9+’ +0 6(0 48)+))0 /)) &)*)* 0)* *)’ +)0 /)) &9)+))* (6) 0) & 0 0 * 0’ 0) &90’ 0 0 ’6) 4 * & 0 + 0 (0 * * / 0 0 0 0 & 0 (9* & 0 ’6/) * (* (0 0 0 09* (* /6/)) () (((& (’ ((() (& ((9(’ (&6/ ((& & (’ (&9& & /6/ + B 78 /)/ / & / &+ / &+ / & / &+ /)/ / & / &+9/)/ / & 6/ / / &+ / &+ /)/ / &+9/)/ / &+6/ / B 78 / &0 / &0 / && / &(/ &) / &) / && / &0 / &(9/ &0 / &)6/ / / &* / &* / &’ / &*9/ &’ / &*6/ / B 78 / ’ / * / * / * / ’ / * / ’ / * / *9/ ’ / * 6/ / / * / * / * / * / *6/ // B: 0 (’ + 0 (9 06/ /& /* /0 /* /09 /* /*6/ / 8B 78 / 0(/ 0 0 / 0* / 0(/ 0) / 0) / 0) / 0 0 / 0(9/ 0* / 0)6/ / / 0& / 0 / 0) / 0 9/ 0) / 0&6/ /(B#")(((&’ &/ &))() (+9)(&+6/ /*)0 &)))) 9)0))6/ /(B#" / ’& / *) / * / ** / ** / ’(/ *+ / ’/ / *’9/ ’& / ’/6/ /(/ *’ / *) / * / *’9/ * / *+6/ / #B4" / 0/ / &+ / 0* / &* / 0) / 0/ / 0(/ 0 0 / &09/ 0* /) 6/ /+ / 0 /) / 0 /) 9/ 0 / 0/6/ / 8B 78 / 0(/ 0 0 / 0* / 0(/ 0) / 0) / 0) / 0 0 / 0(9/ 0* / 0)6/ / / 0& / 0 / 0) / 0 9/ 0) / 0&6/ /(8B4:) +&) () *() &)) +)) ’0) ’&) ’) &)9) () ’&6/ +) +)) ++) 9) +) *06/)/ &B4" / &0 / &* / &) / &) / &+ /)& / & /) / &)9/)& / &’6/ /& / &’ / 0* /)/ / &’9/ 0* /))6/ / with shrubs and vines. Most of the specimens were found on rock boulders, ca. 0.5–1.0 m above the ground, few frogs were in the water. Air temperature was 23.4–29.1o C and relative humidity was 57–79%. Other amphibian species were found at the site, including Rhacophorus kio Ohler & Delorme, Polypedates mutus (Smith), and Kurixalus bisacculus (Taylor). Distribution. Odorrana mutschmanni is currently known only from the type locality in Cao Bang Province, northeastern Vietnam (Fig. 5). This species is presumably also to be found in adjacent karst formations in Ha Giang Province of Vietnam and Guangxi and Yunnan provinces of China. Comparisons. Within the Odorrana andersonii group (O. andersonii, O. jingdongensis, O. kwangwuensis, O. margaretae, O. grahami, O. junlianensis, and O. wuchuanensis), Odorrana mutschmanni sp. nov. differs from O. andersonii by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.5 in males and 0.45 in females in O. andersonii), different ventral color pattern (large dark spots vs. immaculate white in O. andersonii), males without spines on chest (vs. present in O. andersonii), the disc of finger III> 2 times base of phalanges (vs. ≤ 2 times base of phalanges in O. andersonii), and different egg color (wholly unpigmented vs. pigmented in O. andersonii); from O. jingdongensis by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.54 in males and 0.51 in females in O. jingdongensis), different ventral color pattern (large dark spots vs. immaculate white in O. jingdongensis), males without spines on chest (vs. present in O. jingdongensis), and the disc of finger III> 2 times base of phalanges (vs. ≤ 2 times base of phalanges in O. jingdongensis); from O. margaretae by having large dark spots on belly (vs. small dark spots in O. margaretae), males without spines on chest (vs. present in O. margaretae), the disc of finger III> 2 times base of phalanges (vs. ≤ 2 times base of phalanges in O. margaretae), more developed toe webbing (complete to disc on I vs. as narrow fringe to disc on I in O. margaretae), and different egg color (wholly unpigmented vs. pigmented in O. margaretae); from O. kuangwuensis by having a larger body size (SVL 87–92 mm in males and 108–110 mm in females vs. 57 mm in males and 69–71 mm in females in O. kwangwuensis), a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.55 in males and 0.5 in females in O. kwangwuensis), different ventral color pattern (large black spots vs. white with some black spots in O. kwangwuensis), and the disc of finger III> 2 times base of phalanges (vs. ≤ 2 times base of phalanges in O. kwangwuensis); from O. grahami by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.53 in males and 0.48 in females in O. grahami), different ventral color pattern (large black spots vs. immaculate white in O. grahami), males without spines on chest (vs. present in O. grahami), the disc of finger III> 2x base of phalanges (vs. finger III without disc in O. grahami), and different egg color (wholly unpigmented vs. pigmented in O. grahami); from O. junlianensis by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.47 in males and 0.46 in females in O. junlianensis), the absence of external vocal sacs (vs. present in O. junlianensis), and males without spines on chest (vs. white spinules present on the chest in O. junlianensis), the disc of finger III> 2 times base of phalanges (vs. ≤ 2 times base of phalanges in O. junlianensis), and different egg color (wholly unpigmented vs. pigmented in O. junlianensis); from O. wuchuanensis by having a larger size (SVL 87–92 mm in males and 108–110 mm in females vs. 71–77 mm in males and 76–90 mm in females in O. wuchuaensis), a smaller ratio TD/ED (0.70 in males and 0.68 in females vs. 0.83 in males and 0.8 in females in O. wuchuaensis), dorsal surface of head and anterior part of body smooth (vs. shagreened in O. wuchuanensis), different color pattern of flank and limbs (brown vs. green in O. wuchuanensis), and males without white spines on dorsal surface of arm (vs. present in O. wuchuanensis). Odorrana mutschmanni sp. nov. differs from O. absita, O. anlungensis, O. ammaiensis, O. aureola, O. bacboensis, O. banaorum, O. bolavensis, O. chloronota, O. exiliversabilis, O. fengkaiensis, O. gigatympana, O. graminea, O. hainanensis, O. heatwolei, O. hejingensis, O. hosii, O. huanggangensis, O. khalam, O. lipuensis, O. lungshengensis, O. macrotympana, O. monjerai, O. morafkai, O. nanjingensis, O. narina, O. nasica, O. nasuta, O. orba, O. rotodora, O. schmakeri, O. sinica, O. tiananensis, O. tianmuii, O. tormota, O. yentuensis, O. yizhangensis, and O. zhaoi, by having a larger size (SVL 87–92 mm in males vs. ≤ 70 mm in males in other species). Odorrana mutschmanni sp. nov. further differs from O. absita, O. amamiensis, O. aureola, O. banaorum, O. chloronota, O. exiliversabilis, O.gigatympana, O. graminea, O. hosii, O. indeprensa, O. khalam, O. leporipes, O. livida, O. morafkai, O. nasica, O. nasuta, O. orba, O. swinhoana, O. tormota, O. trankieni, O. versabilis, O. yentuensis, and O. zhaoi by the presence of black bars on lips (vs. absent in the latter). Odorrana mutschmanni sp. nov. differs from O. absita, O. alungensis, O. banaorum, O. bolavensis, O. exiliversabilis, O. hosii, O. indeprensa, O. khalam, O. leporpes, O.monjerai, O. narina, O. nasica, O. supranarina, O. tormota, O. trankieni, O. utsunomiyaorum, O. versabilis, O. yentuensis, and O. zhaoi by lacking dorsolateral folds (vs. present in the latter). Odorrana muschmanni sp. nov. differs from O. amamiensis, O. bacboensis, O. bolavensis, O.chapaensis, O. exiliversabilis, O. geminata, O. gigatypana, O. hainanensis, O. heatwolei, O. lipuensis, O. macrotympana, O. nasica, O. orba, O. utsunomiyaorum, O. supranarina, O. tiannanensis, O. tormota, O. versabilis, and O. yentuensis by having a green dorsum (vs. brown, light brown, olive-brown, reddish-brown, gray-blue, iridescent blue or grassgreen in the latter). Odorrana mutschmanni sp. nov. differs from O. absita, O. aureola, O. bacboensis, O. banaorum, O. bolavensis, O. chapaensis, O. chloronota, O. fengkaiensis, O. geminata, O. graminea, O. gigatympana, O. heatwolei, O. indeprensa, O. ishikawae, O. khalam, O. lungshengensis, O. morafkai, O. nasica, O. orba, O. swinhoana, O. tinananensis, O. tormota, O. trankieni, O. yentuensis, O. yizhangensis, and O. zhaoi by the absence of vocal sacs in males (vs. present in the latter)

Odorrana mutschmanni sp. nov. (Figs. 2, 3) Holotype: IEBR 3723 (Field No. CB 2015.12), adult male, collected by T.Q. Nguyen on 22 April 2015 in the karst forest near Coong Village (22o42.712���N, 106o40.075���E, at an elevation of 447 m), Duc Quang Commune, Ha Lang District, Cao Bang Province, Vietnam. Paratypes: IEBR 3724 (Field No. CB 2012.77), adult males, collected on 15 April 2012; IEBR 3725 (Field No. CB 2012.89), adult female and IEBR 3726���3729 (Field No. CB 2012.90���93), adult males, collected on 16 April 2012, by T.Q. Nguyen et al.; ZFMK 97329, 97330 (Field No. CB 2012.139, 2012.140), adult males, collected on 3 May 2012, by H. T. An, S. Herbst and T. Lehmann; IEBR 3730 (Field No. CB2014.16), adult female, collected by C.T. Pham et al. on 10 June 2014, IEBR 3731 (Field No. CB 2015.11), adult female, collected by T.Q. Nguyen on 22 April 2015, the same data as the holotype. Diagnosis. The new species was strongly supported as a member of Odorrana based on molecular analyses (Fig. 1) and is distinguishable from its congeners by a combination of the following morphological characters: (1) size large (SVL 85.9���91.6 mm in males, 108.7���110.1 mm in females); (2) head longer than wide; (3) vomerine teeth present; (4) external vocal sacs absent; (5) snout short (SL/SVL 0.16���0.17); (6) tympanum large (TD/ED 0.70 in males, 0.68 in females); (7) dorsal surface of head and anterior part of body smooth, posterior part of body and flanks with small tubercles; (8) supratympanic fold present; (9) dorsolateral fold absent; (10) webbing formula I0��� 0II0���0III0���1/ 2IV 1/2���0V; (11) in life, dorsum green with dark brown spots; (12) flanks greyish brown with dark brown spots; (13) throat and chest grey, underside of limbs with large dark brown spots, edged in white, forming a network. Description of holotype. Adult male; SVL 85.9 mm; head longer than wide (HL 33.6 mm, HW 29.9 mm); snout round anteriorly in dorsal view, projecting beyond lower jaw; nostril lateral, closer to the snout tip than to eye (NS 6.2 mm, EN 7.5 mm); canthus rostralis distinct; pupil horizontally oval; loreal region slightly concave and oblique; snout length greater than eye diameter (SL 14.0 mm, ED 9.9 mm); internarial distance wider than interorbital distance and upper eyelid (IND 10.5. mm, IOD 9.7 mm, UEW 6.6 mm); tympanum distinct, round, 70% eye diameter (TD 6.9 mm); vomerine teeth in two oblique ridges; tongue cordiform, deeply notched posteriorly; vocal sac absent. Forelimbs: Forelimb length (FLL 16.6 mm), hand length (HAL 44.4 mm); relative finger lengths: II 2 times of the width of phalanges and about 40% the diameter of tympanum; subarticular tubercles round, formula 1, 1, 2, 2; inner metatarsal tubercle oval, elongate; outer metatarsal tubercle small; finger I with nuptial pad, elongate. Hindlimbs: Tibia longer than thigh (FeL 38.1 mm, TbL 46.9 mm), approximately five times longer than wide (TbW 9.8 mm); tips of toes expanded into discs, with circummarginal grooves; width of toe IV disc narrower than width of finger III disc, approximately two times of the width of phalanges; relative length of toes: IVariation. Measurements and morphological characters of the type series are given in Table 3. Sexual dimorphism. The males are smaller than the females in size (SVL: 85.9���91.6 mm, n = 8 vs 108.7��� 110.1 mm, n = 3, respectively). Etymology. We name this new species in honor of Dr. Frank Mutschmann, director of EXOMED in Berlin, in recognition of his support of our research and conservation work in Vietnam. As common names we suggest Mutschmann���s Frog (English), ���ch đ�� mut-x-man (Vietnamese), and Mutschmanns Frosch (German). Ecological notes. Odorrana mutschmanni sp. nov. appears closely associated with karst environments. Specimens were found at night between 19:00 and 23:00 h around a water pool near Coong Village, Duc Quang Commune (Fig. 4). The surrounding habitat was a secondary karst forest of medium and small hardwoods mixed Odorrana mutschmanni !" / & ���(��� () ��� * (��� (��� + ( ��� ( ��� (& ��� && ��� (0 ��� & / & & & & & & & & &��� ��� & .. % % % % % % % % % #! $ ! # $ ! # $ # $ !! $ # #! $ ��,,�� $! # # $! # $! 1 2 2 2 2 2 2 2 2 5 1 6 " 3 3 3 5 1 6 " + + & & 78 +0 * +0 + + / / +* /) * +0 +9 * ++ 6() /+ ��� / * / /+ ���9 / / 06/ ��� 8 && * &* 0 && 0 &) / &) / && &) &* & && 09&* 0 &) 0 6 () () ���)& 0) (9)& 0)(6 (: (& ��� ((/ (+ (* &/ ��� &/ * (+ 9& ��� &/ /6/ &+ ��� & 0) &+ ���9) & +6 (������continued on the next page @ !" / & () (* ��� ��� ((+ (& (&& (0 /& ( ��� & & ��� ��� & & ��� & ��� & ��� ��� & ��� & ��� & & ���.. % % % % % % % % % # $ $ # # $ $ # $ # $ # �� �� $ # # $ # $ !!!!!!,,!!! 1 2 2 2 2 2 2 2 2 5 1 6 " 3 3 3 5 1 6 " + + & & 8 + 0 ��� ���) ���) / ��� & ��� ��� + ��� 9 / ��� +6/ ��� ((((& 0 ((0 (((9(& 0 ((���6/ ��� 8) ��� + ��� (��� 0 + * +) * ��� * ��� (9 * + &6/ 4 * + *) 0 ��� * + * ��� * ���) * 0 0 ���9���) * *6/ 0 ��� + + (+ ��� +9+ (+ /6/ (& (��� (0 (0 (0 (��� & (((+ (09& ((���6/ (& /) * &) & /9) * & ���6/ + 8 &��� /)(& &+ ()(+ &+ &���) &��� * &+ &��� /9)(+ &+ 6(&)) 0)+ 0 0& &)) 090& &)+ +6)) 8)��� +)* +))* +)*)+)) ()*)* +9))��� +6 (0��� ��� 0 0 0 */ / 0 0 09*/ / 0��� ���6(&: 0 / + / + ��� / (/) + 0 9 / + / 6/ 0 + & ((& +9 & (()6/ ��� 8 * / *��� (** + ���/ & *) + *0 ��� *��� *0 / *) +9���/ & *��� 6 +& +&) +��� +&)9+��� +0 6(0 48)+))0 /)) &)*)* 0)* *)��� +)0 /)) &9)+))* (6) 0) & 0 0 * 0��� 0) &90��� 0 0 ���6) 4 * & 0 + 0 (0 * * / 0 0 0 0 & 0 (9* & 0 ���6/) * (* (0 0 0 09* (* /6/)) () (((& (��� ((() (& ((9(��� (&6/ ((& & (��� (&9& & /6/ + B 78 /)/ / & / &+ / &+ / & / &+ /)/ / & / &+9/)/ / & 6/ / / &+ / &+ /)/ / &+9/)/ / &+6/ / B 78 / &0 / &0 / && / &(/ &) / &) / && / &0 / &(9/ &0 / &)6/ / / &* / &* / &��� / &*9/ &��� / &*6/ / B 78 / ��� / * / * / * / ��� / * / ��� / * / *9/ ��� / * 6/ / / * / * / * / * / *6/ // B: 0 (��� + 0 (9 06/ /& /* /0 /* /09 /* /*6/ / 8B 78 / 0(/ 0 0 / 0* / 0(/ 0) / 0) / 0) / 0 0 / 0(9/ 0* / 0)6/ / / 0& / 0 / 0) / 0 9/ 0) / 0&6/ /(B#")(((&��� &/ &))() (+9)(&+6/ /*)0 &)))) 9)0))6/ /(B#" / ���& / *) / * / ** / ** / ���(/ *+ / ���/ / *���9/ ���& / ���/6/ /(/ *��� / *) / * / *���9/ * / *+6/ / #B4" / 0/ / &+ / 0* / &* / 0) / 0/ / 0(/ 0 0 / &09/ 0* /) 6/ /+ / 0 /) / 0 /) 9/ 0 / 0/6/ / 8B 78 / 0(/ 0 0 / 0* / 0(/ 0) / 0) / 0) / 0 0 / 0(9/ 0* / 0)6/ / / 0& / 0 / 0) / 0 9/ 0) / 0&6/ /(8B4:) +&) () *() &)) +)) ���0) ���&) ���) &)9) () ���&6/ +) +)) ++) 9) +) *06/)/ &B4" / &0 / &* / &) / &) / &+ /)& / & /) / &)9/)& / &���6/ /& / &��� / 0* /)/ / &���9/ 0* /))6/ / with shrubs and vines. Most of the specimens were found on rock boulders, ca. 0.5���1.0 m above the ground, few frogs were in the water. Air temperature was 23.4���29.1o C and relative humidity was 57���79%. Other amphibian species were found at the site, including Rhacophorus kio Ohler & Delorme, Polypedates mutus (Smith), and Kurixalus bisacculus (Taylor). Distribution. Odorrana mutschmanni is currently known only from the type locality in Cao Bang Province, northeastern Vietnam (Fig. 5). This species is presumably also to be found in adjacent karst formations in Ha Giang Province of Vietnam and Guangxi and Yunnan provinces of China. Comparisons. Within the Odorrana andersonii group (O. andersonii, O. jingdongensis, O. kwangwuensis, O. margaretae, O. grahami, O. junlianensis, and O. wuchuanensis), Odorrana mutschmanni sp. nov. differs from O. andersonii by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.5 in males and 0.45 in females in O. andersonii), different ventral color pattern (large dark spots vs. immaculate white in O. andersonii), males without spines on chest (vs. present in O. andersonii), the disc of finger III> 2 times base of phalanges (vs. ��� 2 times base of phalanges in O. andersonii), and different egg color (wholly unpigmented vs. pigmented in O. andersonii); from O. jingdongensis by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.54 in males and 0.51 in females in O. jingdongensis), different ventral color pattern (large dark spots vs. immaculate white in O. jingdongensis), males without spines on chest (vs. present in O. jingdongensis), and the disc of finger III> 2 times base of phalanges (vs. ��� 2 times base of phalanges in O. jingdongensis); from O. margaretae by having large dark spots on belly (vs. small dark spots in O. margaretae), males without spines on chest (vs. present in O. margaretae), the disc of finger III> 2 times base of phalanges (vs. ��� 2 times base of phalanges in O. margaretae), more developed toe webbing (complete to disc on I vs. as narrow fringe to disc on I in O. margaretae), and different egg color (wholly unpigmented vs. pigmented in O. margaretae); from O. kuangwuensis by having a larger body size (SVL 87���92 mm in males and 108���110 mm in females vs. 57 mm in males and 69���71 mm in females in O. kwangwuensis), a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.55 in males and 0.5 in females in O. kwangwuensis), different ventral color pattern (large black spots vs. white with some black spots in O. kwangwuensis), and the disc of finger III> 2 times base of phalanges (vs. ��� 2 times base of phalanges in O. kwangwuensis); from O. grahami by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.53 in males and 0.48 in females in O. grahami), different ventral color pattern (large black spots vs. immaculate white in O. grahami), males without spines on chest (vs. present in O. grahami), the disc of finger III> 2x base of phalanges (vs. finger III without disc in O. grahami), and different egg color (wholly unpigmented vs. pigmented in O. grahami); from O. junlianensis by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.47 in males and 0.46 in females in O. junlianensis), the absence of external vocal sacs (vs. present in O. junlianensis), and males without spines on chest (vs. white spinules present on the chest in O. junlianensis), the disc of finger III> 2 times base of phalanges (vs. ��� 2 times base of phalanges in O. junlianensis), and different egg color (wholly unpigmented vs. pigmented in O. junlianensis); from O. wuchuanensis by having a larger size (SVL 87���92 mm in males and 108���110 mm in females vs. 71���77 mm in males and 76���90 mm in females in O. wuchuaensis), a smaller ratio TD/ED (0.70 in males and 0.68 in females vs. 0.83 in males and 0.8 in females in O. wuchuaensis), dorsal surface of head and anterior part of body smooth (vs. shagreened in O. wuchuanensis), different color pattern of flank and limbs (brown vs. green in O. wuchuanensis), and males without white spines on dorsal surface of arm (vs. present in O. wuchuanensis). Odorrana mutschmanni sp. nov. differs from O. absita, O. anlungensis, O. ammaiensis, O. aureola, O. bacboensis, O. banaorum, O. bolavensis, O. chloronota, O. exiliversabilis, O. fengkaiensis, O. gigatympana, O. graminea, O. hainanensis, O. heatwolei, O. hejingensis, O. hosii, O. huanggangensis, O. khalam, O. lipuensis, O. lungshengensis, O. macrotympana, O. monjerai, O. morafkai, O. nanjingensis, O. narina, O. nasica, O. nasuta, O. orba, O. rotodora, O. schmakeri, O. sinica, O. tiananensis, O. tianmuii, O. tormota, O. yentuensis, O. yizhangensis, and O. zhaoi, by having a larger size (SVL 87���92 mm in males vs. ��� 70 mm in males in other species). Odorrana mutschmanni sp. nov. further differs from O. absita, O. amamiensis, O. aureola, O. banaorum, O. chloronota, O. exiliversabilis, O.gigatympana, O. graminea, O. hosii, O. indeprensa, O. khalam, O. leporipes, O. livida, O. morafkai, O. nasica, O. nasuta, O. orba, O. swinhoana, O. tormota, O. trankieni, O. versabilis, O. yentuensis, and O. zhaoi by the presence of black bars on lips (vs. absent in the latter). Odorrana mutschmanni sp. nov. differs from O. absita, O. alungensis, O. banaorum, O. bolavensis, O. exiliversabilis, O. hosii, O. indeprensa, O. khalam, O. leporpes, O.monjerai, O. narina, O. nasica, O. supranarina, O. tormota, O. trankieni, O. utsunomiyaorum, O. versabilis, O. yentuensis, and O. zhaoi by lacking dorsolateral folds (vs. present in the latter). Odorrana muschmanni sp. nov. differs from O. amamiensis, O. bacboensis, O. bolavensis, O.chapaensis, O. exiliversabilis, O. geminata, O. gigatypana, O. hainanensis, O. heatwolei, O. lipuensis, O. macrotympana, O. nasica, O. orba, O. utsunomiyaorum, O. supranarina, O. tiannanensis, O. tormota, O. versabilis, and O. yentuensis by having a green dorsum (vs. brown, light brown, olive-brown, reddish-brown, gray-blue, iridescent blue or grassgreen in the latter). Odorrana mutschmanni sp. nov. differs from O. absita, O. aureola, O. bacboensis, O. banaorum, O. bolavensis, O. chapaensis, O. chloronota, O. fengkaiensis, O. geminata, O. graminea, O. gigatympana, O. heatwolei, O. indeprensa, O. ishikawae, O. khalam, O. lungshengensis, O. morafkai, O. nasica, O. orba, O. swinhoana, O. tinananensis, O. tormota, O. trankieni, O. yentuensis, O. yizhangensis, and O. zhaoi by the absence of vocal sacs in males (vs. present in the latter), Published as part of Pham, Cuong The, Nguyen, Truong Quang, Le, Minh Duc, Bonkowski, Michael & Ziegler, Thomas, 2016, A new species of Odorrana (Amphibia: Anura: Ranidae) from Vietnam, pp. 421-435 in Zootaxa 4084 (3) on pages 423-430, DOI: 10.11646/zootaxa.4084.3.7, http://zenodo.org/record/1052219

Pham, Cuong The, Nguyen, Truong Quang, Le, Minh Duc, Bonkowski, Michael, Ziegler, Thomas (2016): A new species of Odorrana (Amphibia: Anura: Ranidae) from Vietnam. Zootaxa 4084 (3): 421-435, DOI: http://doi.org/10.11646/zootaxa.4084.3.7

Sectonema tropicum sp. nov. urn:lsid:zoobank.org:act: 014CA9CB-57EE-43BB-BBFF-0C1A7EBD522D Figs 1���4; Table 1 Diagnosis The new species is characterized by a 2.56���3.24 mm long body, lip region hardly offset by very weak depression and 19���21 ��m broad, odontostyle 20���21 ��m long at its ventral side and 6.6���7.1 times as long as wide, 730���834 ��m long neck, pharyngeal expansion 403���470 ��m long or occupying 52���59% of total neck length, uterus a simple tube-like structure 150���242 ��m long or 1.2���2.5 times the body diameter, pars refringens vaginae present, V = 48���52, short (32���39 ��m, c = 70���88, c��� = 0.5���0.6) and rounded female tail, male tail similar to that of female (31���40 ��m, c = 73���91, c��� = 0.5���0.6), 91���97 ��m long spicules, and only one weakly developed ventromedian supplement. Etymology The specific epithet refers to the tropical area where the new species was collected. Type material examined Holotype VIETNAM: ♀, in acceptable state of preservation, Northern Vietnam, Cao Bang Province, Pia Oac Natural Reserve, 22�� 36���28������ N, 105�� 52���15������ E, tropical evergreen forest soil associated with Machilus sp. and Dimocarpus sp., deposited in the nematode collection of the University of Ja��n, Spain. Paratypes VIETNAM: 4 ♀♀, 3 ������, in acceptable state of preservation, same data as holotype; 1 ♀, 1 ���, in acceptable state of preservation, same locality, deposited in the nematode collection of the Institute of Ecology and Biological Resources, Hanoi, Vietnam; 1 ���, same locality, used for SEM. Description Adult Moderately slender to slender nematodes of medium to big size, 2.56���3.24 mm long. Body cylindrical, distinctly tapering towards the anterior end, less so towards the posterior end as the caudal region is short and rounded. Habitus curved ventrad after fixation, especially in posterior body region, C-, G- or spiral-shaped. Cuticle 3.0���4.5 ��m thick at anterior region, 5���6 ��m in mid-body and 8���10 ��m on tail; consisting of three layers, especially distinguishable at caudal region: thinner outer layer bearing very fine transverse striation across the entire body, thicker intermediate layer with radial striation, and thin inner layer. Lateral chords 13���21 ��m wide at mid-body, occupying about one-sixth (13���18%) of midbody diameter. Two ventral and two dorsal body pores often present at level of odontostyle-odontophore. Lip region hardly offset from the adjacent body by weak, but perceptible depression, 2.8���3.0 times wider than high and up to one-fourth (17���25%) of body diameter at neck base; lips (under SEM) mostly amalgamated, but their perioral part distinctly separated by the existence of six radial, interlabial, deep incisures delimiting six perceptible liplets; button-like labial papillae, the inner ones located at the margin of the oral field and surrounded by two or three concentric annuli, whereas the outer papillae, located a little behind the inner papillae, are surrounded by only one annulus; cephalic papillae pore- rather than button-like, also surrounded by only one ring-like annulus; oral aperture a dorso-ventral, slightly hexagonal orifice, the lip region hence showing a bi-radial symmetry. Amphid fovea cup-shaped, its opening occupying 9���10 ��m or less than one-half (43���49%) of lip region diameter. Cheilostom nearly cylindrical, without any differentiation. Stomatal protruding structure apparently a reduced odontostyle 6.6���7.1 times longer than wide (see: Remarks), its ventral side 1.0���1.1 times longer than lip region diameter and 0.65���0.83% of body length. Guiding ring simple, plicate, at 0.8���0.9 lip region diameters from the anterior end. Odontophore linear, rod-like, 1.7���1.9 times the odontostyle length, somewhat irregular at its base and with (in lateral view) the ventral side slightly longer than the dorsal one (Fig. 2A). Anterior region of pharynx enlarging very gradually; basal expansion 7.1���10.7 times longer than wide, 3.6���5.6 times as long as body diameter, and occupying 52���59% of total neck length; gland nuclei often obscure, located as follows: DN = 60���62 (n = 4); S 1 N 1 = 75 (n = 2); S 1 N 2 = obscure; S 2 N = obscure. Nerve ring located at 182���216 ��m from anterior end or 25���26% of total neck length. Cardia rounded conoid, 13���17 �� 13���17 ��m; a weak ring-like structure is present surrounding its junction to pharyngeal base. Female Genital system didelphic-amphidelphic, with almost equally and well developed branches, the anterior 320���442 ��m or 12���15% of body length and the posterior 348, 440 ��m (n = 2) or 14, 16% of body length (447, 509 ��m or 15, 17% bearing one uterine egg inside). Reflexed ovaries well developed, often surpassing the sphincter level, the anterior 182���319 ��m, the posterior 207���380 ��m long; oocytes arranged first in two or more rows, then in a single row. Oviduct 124���174 ��m long or 1.1���1.8 times the corresponding body diameter, and consisting of a slender part made of prismatic cells and a moderately developed pars dilatata, with visible lumen and often containing sperm cells inside. Oviduct-uterus junction marked by a sphincter. Uterus a simple, tube-like structure 150���190 ��m long or 1.2���1.7 times the corresponding body diameter [223, 242 ��m (n = 2) long or 2.2, 2.5 times the corresponding body diameter, with one uterine egg inside; and 231, 240 ��m (n = 1) long, with two uterine eggs], always containing abundant sperm cells inside. Vagina extending inwards 52���57 ��m or one-half to foursevenths (48���57%) of body diameter: pars proximalis 33���43 �� 23���28 ��m, with somewhat sigmoid walls surrounded by weak musculature; pars refringens with two trapezoidal pieces measuring (in optical section) 13���16 �� 7���9 ��m and with a combined width of 24���28 ��m; pars distalis 5.5���7.0 ��m long. Vulva a nearly equatorial transverse slit. Prerectum 2.0���2.4, rectum 0.7���1.0 anal body diameters long. Tail short and rounded; two pairs of caudal pores, sublateral and close together. Male Genital system diorchic, with opposite testes. In addition to the ad-cloacal pair, situated at 18���21 ��m from cloacal aperture, one weakly developed ventromedian supplement, lying out the range of retracted spicules and located at 80���131 ��m from the ad-cloacal pair. Spicules distinctly robust and massive, especially in its posterior half, 3.6���4.2 times its maximum width, 1.4���1.6 times the body diameter at level of the cloacal aperture: dorsal contour regularly convex, ventral contour slightly concave, with distinct hump and hollow; curvature 121���124��; head occupying 11���16% of spicule total length, its dorsal side visibly curved at its anterior end and longer than the ventral one, which is shorter and almost straight; median piece 5.4���6.2 times longer than wide, occupying 51���59% of spicule maximum width, reaching the posterior tip; posterior end 7���8 ��m wide. Lateral guiding pieces 23���29 ��m long, 5.3��� 7.1 times longer than wide. Prerectum 3.2���4.6, cloaca 0.9���1.3 times the corresponding body width long. Cloacal aperture, as seen under SEM, a curved anteriad, transverse slit. Tail similar to that of female; caudal pores two pairs, one lateral, another subdorsal. Relationships In having a long stomatal protruding structure (20���21 ��m at its ventral side) and short (c��� up to 1.0) and rounded tail, S. tropicum sp. nov. is morphologically very similar to S. demani Altherr, 1965, S. heynsi Altherr, 1968, S. rotundicauda Goodey, 1951, S. septentrionale Pe��a-Santiago & ��lvarez-Ortega, 2015 and S. sica Clark, 1964, but it can be distinguished from all of them in its smaller general size (body length 2.56���3.24 mm, neck 730���834 ��m long vs more than 5 mm, more than 1000 ��m long, respectively), nearly continuous lip region (vs offset by a more or less deep constriction) and stomatal protruding structure of different nature (a reduced odontostyle vs a mural tooth; see Remarks). Besides, the new species differs from S. demani (see recent re-description by Pe��a-Santiago & ��lvarez-Ortega 2014b) in its less slender (a = 21���36 vs a = 48���56) body, narrower (19���21 vs 27���28 ��m broad) lip region, shorter pharyngeal expansion (occupying 52���59 vs 68���69% of total neck length), shorter female tail (32���39 vs 51���55 ��m, c��� = 0.5���0.6 vs c��� = 0.7���0.8), and male present (vs absent). It differs from S. heynsi (see recent re-description by Pe��a-Santiago & ��lvarez-Ortega 2014b) in its narrower (19���21 vs 28 ��m broad) lip region, circumoral area lacking (vs bearing) cilia- or setae-like structures, shorter pharyngeal expansion (occupying 52���59 vs 68% of total neck length), shorter (32���39 vs 54 ��m, c��� = 0.5���0.6 vs c��� = 0.7) female tail, and male present (vs absent). It differs from S. rotundicauda in its less slender (a = 21���36 vs a = 80) body, narrower (19���21 vs about 28 ��m broad) lip region, shorter (31���40 vs 64 ��m) male tail, fewer (one vs four) ventromedian supplements, and shorter (91���97 vs about 155 ��m) spicules. It differs from S. septentrionale in its less slender (a = 21���36 vs a = 47���64) body, narrower (19���21 vs 25���29 ��m broad) lip region, shorter (occupying 52���59 vs 60���73% of total neck length) pharyngeal expansion, uterus simple (vs tripartite) and shorter (150���242 ��m or 1.2���2.5 times the body diameter vs 370���493 ��m or 2.9���4.3 times the corresponding body diameter), shorter (31���40 vs 40���62 ��m, c��� = 0.5���0.6 vs c��� = 0.6���0.8) tail, fewer (one vs 5���11) ventromedian supplements, and shorter (91���97 vs 100���145 ��m) spicules. Finally, it differs from S. sica in its narrower (19���21 vs about 24 ��m broad) lip region, shorter (occupying 52���59% vs two-thirds of total neck length) pharyngeal expansion, more posterior (V = 48���52 vs V = 40) vulva, shorter (32���39 vs 54 ��m, c��� = 0.5���0.6 vs c��� = 0.7) female tail, and male present (vs absent). Moreover, in having a lip region nearly continuous with the adjacent body, the new species resembles S. mucrodens Siddiqi, 1984 and S. truxum Siddiqi, 1984, but it can be easily distinguished from these by the more anterior position of the stomatal protruding structure (its anterior tip in front of the level of guiding ring, just behind the base of lip region vs its anterior tip distinctly behind the level of guiding ring, far from the lip region base) and pars refringens vaginae well (vs faintly) developed. Moreover, it differs from S. mucrodens by its shorter (730���834 vs 1086���1157 ��m) neck, shorter (occupying 52���59 vs 69% of total neck length) pharyngeal expansion, shorter (32���39 ��m, c��� = 0.5���0.6 vs 44���50 ��m, c��� = 0.7) female tail, and male present (vs absent). It differs from S. truxum in its narrower (19���21 vs 24 ��m broad) lip region, shorter (730���834 vs 837���1070 ��m) neck, more anterior (V = 48���52 vs V = 54���56) vulva, and male present (vs absent). Molecular characterisation Two sequences of the D2���D 3 28S rRNA gene were obtained from one female and one male. Both sequences are very similar (99%) and differ by two nucleotides only. The evolutionary relationships of the new species with representatives of the order Dorylaimida are presented in Figs 5���6. The two S. tropicum sp. nov. sequences are clustered together with other available sequences of Sectonema representatives and form a highly supported clade together with members of the genus Metaporcelaimus, thus confirming previous results (��lvarez-Ortega et al. 2013a, b). Remarks The nature of the stomatal protruding structure in the new species is difficult to interpret and deserves further explanation. The specimens examined bear a reduced odontostyle (comparable to that described in the type species S. ventrale Thone, 1930; see recent description by Pe��a-Santiago & ��lvarez-Ortega 2014a) rather than a mural tooth as the base of its dorsal side is visibly sclerotized and seems to join the dorsal stomatal wall (vs dorsal side not scletorized at the base, somewhat sigmoid and not joining the dorsal stomatal wall). Nonetheless, some doubt persists in this matter., Published as part of Nguyen, Thi Anh Duong, Abolafia, Joaqu��n, Bonkowski, Michael, Pe��a-Santiago, Reyes & ��lvarez-Ortega, Sergio, 2016, Two atypical new species of the genus Sectonema Thorne, 1930 (Nematoda, Dorylaimida, Aporcelaimidae) from Vietnam, pp. 1-20 in European Journal of Taxonomy 171 on pages 4-11, DOI: 10.5852/ejt.2016.171, http://zenodo.org/record/3832734, {"references":["Alvarez-Ortega S., Nguyen T. A. D., Abolafia J., Vu T. T. T. & Pena-Santiago R. 2015. Three new species of the genus Aporcelaimoides Heyns, 1965 from Vietnam (Nematoda, Dorylaimida, Aporcelaimidae), and an updated taxonomy of the genus. ZooKeys 516: 1 - 26. http: // dx. doi. org / 10.3897 / zookeys. 516.10087","Pena-Santiago R. & Alvarez-Ortega S. 2014 b. Re-description of three species of the genus Sectonema Thorne, 1930 (Nematoda: Dorylaimida: Aporcelaimidae) originally studied by E. Altherr. Zootaxa 3881: 63 - 74. http: // dx. doi. org / 10.11646 / zootaxa. 3881.1.5","Alvarez-Ortega S., Subbotin S. A. & Pena-Santiago R. 2013 a. Morphological and molecular characterization of Californian species of the genus Metaporcelaimus Lordello, 1965 (Dorylaimida, Aporcelaimidae), with a new concept of the genus. Nematology 15 (3): 251 - 278. http: // dx. doi. org / 10.1163 / 15685411 - 00002674","Pena-Santiago R. & Alvarez-Ortega S. 2014 a. Studies on the genus Sectonema Thorne, 1930 (Dorylaimida: Aporcelaimidae). Re-description of S. ventrale, the type species of the genus. Nematology 16 (9): 1097 - 1104. http: // dx. doi. org / 10.1163 / 15685411 - 00002837"]}

Sectonema vietnamense sp. nov. urn:lsid:zoobank.org:act: AF58451E-D7A7-4E49-8A18-9F3998EF4A87 Figs 7–9, Table 2 Diagnosis The new species is characterized by a slender (a = 33–49) 2.71–4.25 mm long body, lip region offset by depression and 14–16 μm broad, odontostyle 8–9 μm long at its ventral side, 716–918 μm long neck, pharyngeal expansion 463–618 μm long or 63–67% of total neck length, uterus simple and 209–242 μm long or 2.5–2.9 times the corresponding body diameter, pars refringens vaginae absent, V = 54, short (36 μm, c = 105, c’ = 0.7) and rounded tail in female and slightly more conoid (34–39 μm, c = 70– 115, c’ = 0.6–0.8) in male, 59–75 μm long spicules, and three or four irregularly spaced ventromedian supplements bearing hiatus. Etymology The specific epithet refers to the geographical origin of the species. Type material examined Holotype VIETNAM: ♀, in good state of preservation, Northern Vietnam, Cuc Phuong National Park, pristine tropical forest, deposited in the nematode collection of the University of Jaén, Spain. Paratypes VIETNAM: 4 ♁♁, 3 juvs, in good state of preservation, same data as holotype; 1 ♁, in good state of preservation, same locality, deposited in the nematode collection of the Institute of Ecology and Biological Resources, Hanoi, Vietnam; 1♁, 1 juv., same locality, used for SEM. Other material examined VIETNAM: 2 ♁♁, in acceptable state of preservation, Northern Vietnam, Thái Bình province, intensively cultured field, deposited in the nematode collection of the University of Jaén, Spain. Description (type population) Adult Very slender nematodes of big size, 3.13–4.25 mm long. Body cylindrical, distinctly tapering towards the anterior end, less so towards the posterior one as the caudal region is rounded. Habitus regularly curved ventrad after fixation, to a more or less open C, occasionally more curved at posterior body region. Cuticle 2.0–2.5 μm thick at anterior region, 3.0–4.5 μm in mid-body and 8–10 μm on tail; consisting of three layers especially distinguishable at caudal region: thinner outer layer bearing very fine transverse striation through the entire body, thicker intermediate layer with radial striation, and thin inner layer. Lateral chords 6–12 μm wide at mid-body, occupying up to one-sixth (8–15%) of mid-body diameter. Two ventral and two dorsal body pores often present at level of stoma, and four distinct lateral pores 6–20 μm apart visible behind the amphid fovea. Lip region nearly continuous (a shallow depression is, however, often perceptible), weakly angular, visibly narrower than adjacent body, 2.1–2.3 times wider than high and about one-fifth (18–23%) of body diameter at neck base; it appears (under SEM) marked by radial (oral field) or longitudinal (remaining lip region) incisures, with amalgamated lips and protruding, pore-like labial and cephalic papillae, all of them surrounded by a ring-like annulus, and the inner labial papillae visibly larger than the outer ones; oral field perceptibly hexagonal, oral aperture a dorso-ventral slit, the lip region hence showing a bi-radial symmetry. Cupshaped amphid fovea, its opening at level of cephalic depression and occupying 10–11 μm or 68–72% of lip region diameter. Nearly cylindrical cheilostom, without differentiation. Odontostyle comparatively short and occupying the whole stomatal lumen, its ventral side 0.5–0.6 times longer than lip region diameter and 0.20–0.29% of body length. Guiding ring simple, somewhat plicate, at 0.5 times the lip region diameter from anterior end. Odontophore linear, rod-like, 3.7–4.2 times the odontostyle length. Tripartite pharynx, consisting of an anteriorly thickened section behind the odontophore base, a perceptible narrower intermediate section with the nerve ring surrounding it, which enlarges very gradually into the basal expansion 9.9–15.0 times longer than wide, 6.7–7.3 times longer than body diameter and occupying 63–67% of total neck length; gland nuclei mostly obscure, only S 2 N are visible, at 77–78% (n = 3) of total neck length. Nerve ring located at 147–178 μm from anterior end or 19–22% of total neck length. Cardia rounded conoid, 15–19 × 12–17 μm; a ring-like structure is present surrounding its junction to pharyngeal base. Female Genital system didelphic-amphidelphic, with almost equally and well developed branches, the anterior 473 μm long or 13% of body length and the posterior 387 μm long or 10% of body length, each branch bearing one uterine egg. Moderately sized ovaries, not surpassing the sphincter level, the anterior 191 μm, the posterior 141 μm long; oocytes arranged first in two or more rows, then in a single row. Oviduct 117, 142 μm long or 1.4, 1.7 times the corresponding body diameter, and consisting of a slender part made of prismatic cells and a well developed pars dilatata with wide lumen and sperm cells inside. Oviduct-uterus junction marked by a sphincter. Uterus a simple, tube-like structure 209, 242 μm long or 2.5, 2.9 times the corresponding body diameter, containing abundant sperm cells and one uterine egg. Vagina extending inwards 36 μm or about three-sevenths (44%) of body diameter: pars proximalis 23 × 28 μm, somewhat sigmoid walls and surrounded by weak musculature; pars refringens absent; pars distalis well developed, 15 μm long. Vulva a post-equatorial transverse slit. Prerectum 3.0, rectum 0.9 anal body diameters long. Short and rounded tail, its inner core with a finger-like projection at tail end; two pairs of caudal pores, one sub-lateral, another sub-dorsal. Male Genital system diorchic, with opposite testes. In addition to the ad-cloacal pair, situated at 14–19 μm from cloacal aperture, there is a series of three or four irregularly spaced, 15–48 μm apart, ventromedian supplements, the posteriormost of which lying out the range of retracted spicules, at 76–91 μm from the ad-cloacal pair. Spicule robust and massive, especially in its posterior half, 3.6–4.6 times its maximum width, 1.3–1.4 times the body diameter at level of the cloacal aperture: dorsal contour regularly convex, ventral contour slightly concave, with distinct hump and hollow; curvature 125–136º; head occupying 13–19% of spicule total length, its dorsal contour distinct curved at its anterior end and longer than the ventral one, which is short and almost straight; median piece 5.9–7.1 times longer than wide, occupying 52–65% of spicule maximum width, reaching the posterior tip; posterior end 6–7 μm wide. Lateral guiding pieces 15–17 μm long, 5.3–6.5 times longer than wide. Prerectum 4.3–5.7, cloaca 1.0–1.3 times the corresponding body width long. Rectal glands and their ducts (cf. Coomans & Loof 1986) very well perceptible (Fig. 8K). Cloacal aperture, as seen under SEM, a somewhat curved anteriad, transverse slit; the two supplements of the precloacal pair distinctly separated. Tail a little more conoid than that of female; two pairs of caudal pores, one almost lateral, another sub-dorsal. Relationships The new species is easily distinguishable from its congeners in the unique combination of these features: narrow and nearly continuous lip region, very short odontostyle, absence of pars refringens vaginae and short and rounded tail. Nonetheless, it resembles S. barbatum Heyns, 1965 in its general morphology, but it can be distinguished from this because of its oral field lacking any setae-like structure (vs bearing numerous short forwardly pointing setae), narrower (14–16 vs about 22 μm broad) lip region, shorter (8–9 vs 16 μm) odontostyle, pars refringens vaginae absent (vs present), shorter (36 μm, c = 105, c’ = 0.7 vs 48 μm, c = 81, c’ = 1.0) female tail, and male present (vs absent). In having a comparatively small body size and nearly continuous lip region, S. vietnamense sp. nov. is close to S. mucrodens and S. truxum, from which it differs in its narrower (14–16 vs more than 20 μm broad) lip region, shorter stomatal protruding structure (8–9 vs more than 20 μm) located as usual (vs the anterior tip distinctly behind the level of guiding ring), and pars refringens vaginae absent (vs consisting of two small pieces). Sectonema vietnamense sp. nov. is also similar to S. pseudoventrale Heyns, 1965 in having a short odontostyle, but can be distinguished from this by its narrower (14–16 vs 21–23 μm broad) lip region, shorter (209–242 μm or 2.5–2.9 times the corresponding body diameter vs 123 μm or 1.3 times the corresponding body diameter) uterus, shorter (36 μm, c = 105, c’ = 0.7 vs 46–63 μm, c = 82–130, c’ = 0.8–1.0) female tail, shorter (59– 75 vs 90–95 μm) spicules, and fewer (three or four vs six or seven) ventromedian supplements. Finally, the new species also resembles S. tropicum sp. nov., but it can be distinguished from this by its narrower (14–16 vs 19–21 μm broad) lip region, shorter (8–9 vs 20–21 μm at its ventral side) odontostyle, longer (occupying 63–67 vs 52–59% of total neck length) pharyngeal expansion, more posterior vulva (V = 54 vs V =48–52), pars refringens vaginae absent (vs present), shorter (59–75 vs 91–97 μm) spicules and higher number of ventromedian supplements (three or four vs one). Remarks The two males from Thái Bình are very similar to those of the type population, but some minor morphometric differences have been also noted: somewhat smaller (2.71, 3.02 vs 3.13–4.25 mm long) and more obese (a = 33, 35 vs a = 41–49) body. Such differences have been regarded as instraspecific variation as the number of specimens examined is low and the main morphological and morphometric features are coincident.

Sectonema vietnamense sp. nov. urn:lsid:zoobank.org:act: AF58451E-D7A7-4E49-8A18-9F3998EF4A87 Figs 7���9, Table 2 Diagnosis The new species is characterized by a slender (a = 33���49) 2.71���4.25 mm long body, lip region offset by depression and 14���16 ��m broad, odontostyle 8���9 ��m long at its ventral side, 716���918 ��m long neck, pharyngeal expansion 463���618 ��m long or 63���67% of total neck length, uterus simple and 209���242 ��m long or 2.5���2.9 times the corresponding body diameter, pars refringens vaginae absent, V = 54, short (36 ��m, c = 105, c��� = 0.7) and rounded tail in female and slightly more conoid (34���39 ��m, c = 70��� 115, c��� = 0.6���0.8) in male, 59���75 ��m long spicules, and three or four irregularly spaced ventromedian supplements bearing hiatus. Etymology The specific epithet refers to the geographical origin of the species. Type material examined Holotype VIETNAM: ♀, in good state of preservation, Northern Vietnam, Cuc Phuong National Park, pristine tropical forest, deposited in the nematode collection of the University of Ja��n, Spain. Paratypes VIETNAM: 4 ������, 3 juvs, in good state of preservation, same data as holotype; 1 ���, in good state of preservation, same locality, deposited in the nematode collection of the Institute of Ecology and Biological Resources, Hanoi, Vietnam; 1���, 1 juv., same locality, used for SEM. Other material examined VIETNAM: 2 ������, in acceptable state of preservation, Northern Vietnam, Th��i B��nh province, intensively cultured field, deposited in the nematode collection of the University of Ja��n, Spain. Description (type population) Adult Very slender nematodes of big size, 3.13���4.25 mm long. Body cylindrical, distinctly tapering towards the anterior end, less so towards the posterior one as the caudal region is rounded. Habitus regularly curved ventrad after fixation, to a more or less open C, occasionally more curved at posterior body region. Cuticle 2.0���2.5 ��m thick at anterior region, 3.0���4.5 ��m in mid-body and 8���10 ��m on tail; consisting of three layers especially distinguishable at caudal region: thinner outer layer bearing very fine transverse striation through the entire body, thicker intermediate layer with radial striation, and thin inner layer. Lateral chords 6���12 ��m wide at mid-body, occupying up to one-sixth (8���15%) of mid-body diameter. Two ventral and two dorsal body pores often present at level of stoma, and four distinct lateral pores 6���20 ��m apart visible behind the amphid fovea. Lip region nearly continuous (a shallow depression is, however, often perceptible), weakly angular, visibly narrower than adjacent body, 2.1���2.3 times wider than high and about one-fifth (18���23%) of body diameter at neck base; it appears (under SEM) marked by radial (oral field) or longitudinal (remaining lip region) incisures, with amalgamated lips and protruding, pore-like labial and cephalic papillae, all of them surrounded by a ring-like annulus, and the inner labial papillae visibly larger than the outer ones; oral field perceptibly hexagonal, oral aperture a dorso-ventral slit, the lip region hence showing a bi-radial symmetry. Cupshaped amphid fovea, its opening at level of cephalic depression and occupying 10���11 ��m or 68���72% of lip region diameter. Nearly cylindrical cheilostom, without differentiation. Odontostyle comparatively short and occupying the whole stomatal lumen, its ventral side 0.5���0.6 times longer than lip region diameter and 0.20���0.29% of body length. Guiding ring simple, somewhat plicate, at 0.5 times the lip region diameter from anterior end. Odontophore linear, rod-like, 3.7���4.2 times the odontostyle length. Tripartite pharynx, consisting of an anteriorly thickened section behind the odontophore base, a perceptible narrower intermediate section with the nerve ring surrounding it, which enlarges very gradually into the basal expansion 9.9���15.0 times longer than wide, 6.7���7.3 times longer than body diameter and occupying 63���67% of total neck length; gland nuclei mostly obscure, only S 2 N are visible, at 77���78% (n = 3) of total neck length. Nerve ring located at 147���178 ��m from anterior end or 19���22% of total neck length. Cardia rounded conoid, 15���19 �� 12���17 ��m; a ring-like structure is present surrounding its junction to pharyngeal base. Female Genital system didelphic-amphidelphic, with almost equally and well developed branches, the anterior 473 ��m long or 13% of body length and the posterior 387 ��m long or 10% of body length, each branch bearing one uterine egg. Moderately sized ovaries, not surpassing the sphincter level, the anterior 191 ��m, the posterior 141 ��m long; oocytes arranged first in two or more rows, then in a single row. Oviduct 117, 142 ��m long or 1.4, 1.7 times the corresponding body diameter, and consisting of a slender part made of prismatic cells and a well developed pars dilatata with wide lumen and sperm cells inside. Oviduct-uterus junction marked by a sphincter. Uterus a simple, tube-like structure 209, 242 ��m long or 2.5, 2.9 times the corresponding body diameter, containing abundant sperm cells and one uterine egg. Vagina extending inwards 36 ��m or about three-sevenths (44%) of body diameter: pars proximalis 23 �� 28 ��m, somewhat sigmoid walls and surrounded by weak musculature; pars refringens absent; pars distalis well developed, 15 ��m long. Vulva a post-equatorial transverse slit. Prerectum 3.0, rectum 0.9 anal body diameters long. Short and rounded tail, its inner core with a finger-like projection at tail end; two pairs of caudal pores, one sub-lateral, another sub-dorsal. Male Genital system diorchic, with opposite testes. In addition to the ad-cloacal pair, situated at 14���19 ��m from cloacal aperture, there is a series of three or four irregularly spaced, 15���48 ��m apart, ventromedian supplements, the posteriormost of which lying out the range of retracted spicules, at 76���91 ��m from the ad-cloacal pair. Spicule robust and massive, especially in its posterior half, 3.6���4.6 times its maximum width, 1.3���1.4 times the body diameter at level of the cloacal aperture: dorsal contour regularly convex, ventral contour slightly concave, with distinct hump and hollow; curvature 125���136��; head occupying 13���19% of spicule total length, its dorsal contour distinct curved at its anterior end and longer than the ventral one, which is short and almost straight; median piece 5.9���7.1 times longer than wide, occupying 52���65% of spicule maximum width, reaching the posterior tip; posterior end 6���7 ��m wide. Lateral guiding pieces 15���17 ��m long, 5.3���6.5 times longer than wide. Prerectum 4.3���5.7, cloaca 1.0���1.3 times the corresponding body width long. Rectal glands and their ducts (cf. Coomans & Loof 1986) very well perceptible (Fig. 8K). Cloacal aperture, as seen under SEM, a somewhat curved anteriad, transverse slit; the two supplements of the precloacal pair distinctly separated. Tail a little more conoid than that of female; two pairs of caudal pores, one almost lateral, another sub-dorsal. Relationships The new species is easily distinguishable from its congeners in the unique combination of these features: narrow and nearly continuous lip region, very short odontostyle, absence of pars refringens vaginae and short and rounded tail. Nonetheless, it resembles S. barbatum Heyns, 1965 in its general morphology, but it can be distinguished from this because of its oral field lacking any setae-like structure (vs bearing numerous short forwardly pointing setae), narrower (14���16 vs about 22 ��m broad) lip region, shorter (8���9 vs 16 ��m) odontostyle, pars refringens vaginae absent (vs present), shorter (36 ��m, c = 105, c��� = 0.7 vs 48 ��m, c = 81, c��� = 1.0) female tail, and male present (vs absent). In having a comparatively small body size and nearly continuous lip region, S. vietnamense sp. nov. is close to S. mucrodens and S. truxum, from which it differs in its narrower (14���16 vs more than 20 ��m broad) lip region, shorter stomatal protruding structure (8���9 vs more than 20 ��m) located as usual (vs the anterior tip distinctly behind the level of guiding ring), and pars refringens vaginae absent (vs consisting of two small pieces). Sectonema vietnamense sp. nov. is also similar to S. pseudoventrale Heyns, 1965 in having a short odontostyle, but can be distinguished from this by its narrower (14���16 vs 21���23 ��m broad) lip region, shorter (209���242 ��m or 2.5���2.9 times the corresponding body diameter vs 123 ��m or 1.3 times the corresponding body diameter) uterus, shorter (36 ��m, c = 105, c��� = 0.7 vs 46���63 ��m, c = 82���130, c��� = 0.8���1.0) female tail, shorter (59��� 75 vs 90���95 ��m) spicules, and fewer (three or four vs six or seven) ventromedian supplements. Finally, the new species also resembles S. tropicum sp. nov., but it can be distinguished from this by its narrower (14���16 vs 19���21 ��m broad) lip region, shorter (8���9 vs 20���21 ��m at its ventral side) odontostyle, longer (occupying 63���67 vs 52���59% of total neck length) pharyngeal expansion, more posterior vulva (V = 54 vs V =48���52), pars refringens vaginae absent (vs present), shorter (59���75 vs 91���97 ��m) spicules and higher number of ventromedian supplements (three or four vs one). Remarks The two males from Th��i B��nh are very similar to those of the type population, but some minor morphometric differences have been also noted: somewhat smaller (2.71, 3.02 vs 3.13���4.25 mm long) and more obese (a = 33, 35 vs a = 41���49) body. Such differences have been regarded as instraspecific variation as the number of specimens examined is low and the main morphological and morphometric features are coincident., Published as part of Nguyen, Thi Anh Duong, Abolafia, Joaqu��n, Bonkowski, Michael, Pe��a-Santiago, Reyes & ��lvarez-Ortega, Sergio, 2016, Two atypical new species of the genus Sectonema Thorne, 1930 (Nematoda, Dorylaimida, Aporcelaimidae) from Vietnam, pp. 1-20 in European Journal of Taxonomy 171 on pages 13-18, DOI: 10.5852/ejt.2016.171, http://zenodo.org/record/3832734, {"references":["Coomans A. & Loof P. A. A. 1986. Observations on the glands of the male reproductive system in dorylaims and its phylogenetic importance. Revue de Nematologie 9 (3): 261 - 265."]}

Sectonema tropicum sp. nov. urn:lsid:zoobank.org:act: 014CA9CB-57EE-43BB-BBFF-0C1A7EBD522D Figs 1–4; Table 1 Diagnosis The new species is characterized by a 2.56–3.24 mm long body, lip region hardly offset by very weak depression and 19–21 μm broad, odontostyle 20–21 μm long at its ventral side and 6.6–7.1 times as long as wide, 730–834 μm long neck, pharyngeal expansion 403–470 μm long or occupying 52–59% of total neck length, uterus a simple tube-like structure 150–242 μm long or 1.2–2.5 times the body diameter, pars refringens vaginae present, V = 48–52, short (32–39 μm, c = 70–88, c’ = 0.5–0.6) and rounded female tail, male tail similar to that of female (31–40 μm, c = 73–91, c’ = 0.5–0.6), 91–97 μm long spicules, and only one weakly developed ventromedian supplement. Etymology The specific epithet refers to the tropical area where the new species was collected. Type material examined Holotype VIETNAM: ♀, in acceptable state of preservation, Northern Vietnam, Cao Bang Province, Pia Oac Natural Reserve, 22º 36’28’’ N, 105º 52’15’’ E, tropical evergreen forest soil associated with Machilus sp. and Dimocarpus sp., deposited in the nematode collection of the University of Jaén, Spain. Paratypes VIETNAM: 4 ♀♀, 3 ♁♁, in acceptable state of preservation, same data as holotype; 1 ♀, 1 ♁, in acceptable state of preservation, same locality, deposited in the nematode collection of the Institute of Ecology and Biological Resources, Hanoi, Vietnam; 1 ♁, same locality, used for SEM. Description Adult Moderately slender to slender nematodes of medium to big size, 2.56–3.24 mm long. Body cylindrical, distinctly tapering towards the anterior end, less so towards the posterior end as the caudal region is short and rounded. Habitus curved ventrad after fixation, especially in posterior body region, C-, G- or spiral-shaped. Cuticle 3.0–4.5 μm thick at anterior region, 5–6 μm in mid-body and 8–10 μm on tail; consisting of three layers, especially distinguishable at caudal region: thinner outer layer bearing very fine transverse striation across the entire body, thicker intermediate layer with radial striation, and thin inner layer. Lateral chords 13–21 μm wide at mid-body, occupying about one-sixth (13–18%) of midbody diameter. Two ventral and two dorsal body pores often present at level of odontostyle-odontophore. Lip region hardly offset from the adjacent body by weak, but perceptible depression, 2.8–3.0 times wider than high and up to one-fourth (17–25%) of body diameter at neck base; lips (under SEM) mostly amalgamated, but their perioral part distinctly separated by the existence of six radial, interlabial, deep incisures delimiting six perceptible liplets; button-like labial papillae, the inner ones located at the margin of the oral field and surrounded by two or three concentric annuli, whereas the outer papillae, located a little behind the inner papillae, are surrounded by only one annulus; cephalic papillae pore- rather than button-like, also surrounded by only one ring-like annulus; oral aperture a dorso-ventral, slightly hexagonal orifice, the lip region hence showing a bi-radial symmetry. Amphid fovea cup-shaped, its opening occupying 9–10 μm or less than one-half (43–49%) of lip region diameter. Cheilostom nearly cylindrical, without any differentiation. Stomatal protruding structure apparently a reduced odontostyle 6.6–7.1 times longer than wide (see: Remarks), its ventral side 1.0–1.1 times longer than lip region diameter and 0.65–0.83% of body length. Guiding ring simple, plicate, at 0.8–0.9 lip region diameters from the anterior end. Odontophore linear, rod-like, 1.7–1.9 times the odontostyle length, somewhat irregular at its base and with (in lateral view) the ventral side slightly longer than the dorsal one (Fig. 2A). Anterior region of pharynx enlarging very gradually; basal expansion 7.1–10.7 times longer than wide, 3.6–5.6 times as long as body diameter, and occupying 52–59% of total neck length; gland nuclei often obscure, located as follows: DN = 60–62 (n = 4); S 1 N 1 = 75 (n = 2); S 1 N 2 = obscure; S 2 N = obscure. Nerve ring located at 182–216 μm from anterior end or 25–26% of total neck length. Cardia rounded conoid, 13–17 × 13–17 μm; a weak ring-like structure is present surrounding its junction to pharyngeal base. Female Genital system didelphic-amphidelphic, with almost equally and well developed branches, the anterior 320–442 μm or 12–15% of body length and the posterior 348, 440 μm (n = 2) or 14, 16% of body length (447, 509 μm or 15, 17% bearing one uterine egg inside). Reflexed ovaries well developed, often surpassing the sphincter level, the anterior 182–319 μm, the posterior 207–380 μm long; oocytes arranged first in two or more rows, then in a single row. Oviduct 124–174 μm long or 1.1–1.8 times the corresponding body diameter, and consisting of a slender part made of prismatic cells and a moderately developed pars dilatata, with visible lumen and often containing sperm cells inside. Oviduct-uterus junction marked by a sphincter. Uterus a simple, tube-like structure 150–190 μm long or 1.2–1.7 times the corresponding body diameter [223, 242 μm (n = 2) long or 2.2, 2.5 times the corresponding body diameter, with one uterine egg inside; and 231, 240 μm (n = 1) long, with two uterine eggs], always containing abundant sperm cells inside. Vagina extending inwards 52–57 μm or one-half to foursevenths (48–57%) of body diameter: pars proximalis 33–43 × 23–28 μm, with somewhat sigmoid walls surrounded by weak musculature; pars refringens with two trapezoidal pieces measuring (in optical section) 13–16 × 7–9 μm and with a combined width of 24–28 μm; pars distalis 5.5–7.0 μm long. Vulva a nearly equatorial transverse slit. Prerectum 2.0–2.4, rectum 0.7–1.0 anal body diameters long. Tail short and rounded; two pairs of caudal pores, sublateral and close together. Male Genital system diorchic, with opposite testes. In addition to the ad-cloacal pair, situated at 18–21 μm from cloacal aperture, one weakly developed ventromedian supplement, lying out the range of retracted spicules and located at 80–131 μm from the ad-cloacal pair. Spicules distinctly robust and massive, especially in its posterior half, 3.6–4.2 times its maximum width, 1.4–1.6 times the body diameter at level of the cloacal aperture: dorsal contour regularly convex, ventral contour slightly concave, with distinct hump and hollow; curvature 121–124°; head occupying 11–16% of spicule total length, its dorsal side visibly curved at its anterior end and longer than the ventral one, which is shorter and almost straight; median piece 5.4–6.2 times longer than wide, occupying 51–59% of spicule maximum width, reaching the posterior tip; posterior end 7–8 μm wide. Lateral guiding pieces 23–29 μm long, 5.3– 7.1 times longer than wide. Prerectum 3.2–4.6, cloaca 0.9–1.3 times the corresponding body width long. Cloacal aperture, as seen under SEM, a curved anteriad, transverse slit. Tail similar to that of female; caudal pores two pairs, one lateral, another subdorsal. Relationships In having a long stomatal protruding structure (20–21 μm at its ventral side) and short (c’ up to 1.0) and rounded tail, S. tropicum sp. nov. is morphologically very similar to S. demani Altherr, 1965, S. heynsi Altherr, 1968, S. rotundicauda Goodey, 1951, S. septentrionale Peña-Santiago & Álvarez-Ortega, 2015 and S. sica Clark, 1964, but it can be distinguished from all of them in its smaller general size (body length 2.56–3.24 mm, neck 730–834 μm long vs more than 5 mm, more than 1000 μm long, respectively), nearly continuous lip region (vs offset by a more or less deep constriction) and stomatal protruding structure of different nature (a reduced odontostyle vs a mural tooth; see Remarks). Besides, the new species differs from S. demani (see recent re-description by Peña-Santiago & Álvarez-Ortega 2014b) in its less slender (a = 21–36 vs a = 48–56) body, narrower (19–21 vs 27–28 μm broad) lip region, shorter pharyngeal expansion (occupying 52–59 vs 68–69% of total neck length), shorter female tail (32–39 vs 51–55 μm, c’ = 0.5–0.6 vs c’ = 0.7–0.8), and male present (vs absent). It differs from S. heynsi (see recent re-description by Peña-Santiago & Álvarez-Ortega 2014b) in its narrower (19–21 vs 28 μm broad) lip region, circumoral area lacking (vs bearing) cilia- or setae-like structures, shorter pharyngeal expansion (occupying 52–59 vs 68% of total neck length), shorter (32–39 vs 54 μm, c’ = 0.5–0.6 vs c’ = 0.7) female tail, and male present (vs absent). It differs from S. rotundicauda in its less slender (a = 21–36 vs a = 80) body, narrower (19–21 vs about 28 μm broad) lip region, shorter (31–40 vs 64 μm) male tail, fewer (one vs four) ventromedian supplements, and shorter (91–97 vs about 155 μm) spicules. It differs from S. septentrionale in its less slender (a = 21–36 vs a = 47–64) body, narrower (19–21 vs 25–29 μm broad) lip region, shorter (occupying 52–59 vs 60–73% of total neck length) pharyngeal expansion, uterus simple (vs tripartite) and shorter (150–242 μm or 1.2–2.5 times the body diameter vs 370–493 μm or 2.9–4.3 times the corresponding body diameter), shorter (31–40 vs 40–62 μm, c’ = 0.5–0.6 vs c’ = 0.6–0.8) tail, fewer (one vs 5–11) ventromedian supplements, and shorter (91–97 vs 100–145 μm) spicules. Finally, it differs from S. sica in its narrower (19–21 vs about 24 μm broad) lip region, shorter (occupying 52–59% vs two-thirds of total neck length) pharyngeal expansion, more posterior (V = 48–52 vs V = 40) vulva, shorter (32–39 vs 54 μm, c’ = 0.5–0.6 vs c’ = 0.7) female tail, and male present (vs absent). Moreover, in having a lip region nearly continuous with the adjacent body, the new species resembles S. mucrodens Siddiqi, 1984 and S. truxum Siddiqi, 1984, but it can be easily distinguished from these by the more anterior position of the stomatal protruding structure (its anterior tip in front of the level of guiding ring, just behind the base of lip region vs its anterior tip distinctly behind the level of guiding ring, far from the lip region base) and pars refringens vaginae well (vs faintly) developed. Moreover, it differs from S. mucrodens by its shorter (730–834 vs 1086–1157 μm) neck, shorter (occupying 52–59 vs 69% of total neck length) pharyngeal expansion, shorter (32–39 μm, c’ = 0.5–0.6 vs 44–50 μm, c’ = 0.7) female tail, and male present (vs absent). It differs from S. truxum in its narrower (19–21 vs 24 μm broad) lip region, shorter (730–834 vs 837–1070 μm) neck, more anterior (V = 48–52 vs V = 54–56) vulva, and male present (vs absent). Molecular characterisation Two sequences of the D2–D 3 28S rRNA gene were obtained from one female and one male. Both sequences are very similar (99%) and differ by two nucleotides only. The evolutionary relationships of the new species with representatives of the order Dorylaimida are presented in Figs 5–6. The two S. tropicum sp. nov. sequences are clustered together with other available sequences of Sectonema representatives and form a highly supported clade together with members of the genus Metaporcelaimus, thus confirming previous results (Álvarez-Ortega et al. 2013a, b). Remarks The nature of the stomatal protruding structure in the new species is difficult to interpret and deserves further explanation. The specimens examined bear a reduced odontostyle (comparable to that described in the type species S. ventrale Thone, 1930; see recent description by Peña-Santiago & Álvarez-Ortega 2014a) rather than a mural tooth as the base of its dorsal side is visibly sclerotized and seems to join the dorsal stomatal wall (vs dorsal side not scletorized at the base, somewhat sigmoid and not joining the dorsal stomatal wall). Nonetheless, some doubt persists in this matter.

Cyrtodactylus soudthichaki sp. nov. (Figs. 1–3) Holotype. VFU R. 2015.18, adult male, Khun Don region (17 o 33.731 ’N, 104 o 52.360 ’E, elevation 167 m a.s.l.) within Phou Hin Poun NPA, Khammouane Province, central Laos, collected during night time between 19:00 and 21:00 on 4 April 2015 by V. Q. Luu and T. Calame. Paratypes. IEBR A. 2015.34, adult female and NUOL R- 2015.5, subadult male, bearing the same data as the holotype. Diagnosis. The new species can be distinguished from all other members of the genus Cyrtodactylus from the mainland Indochina region by a combination of the following characters: adult SVL 69.2 –70.0 mm; dorsal head and neck with dark blotches; nuchal loop present; dorsum with five brown bands between limb insertions; 19 or 20 irregular rows of dorsal tubercles; 32 or 33 ventral scale rows; ventrolateral folds present with distinct tubercles; dorsal surface of hind limbs with tubercles; 29 precloacal and femoral pores in a continuous row in males, precloacal pores absent in the female; enlarged femoral and precloacal scales present; 4 or 5 postcloacal tubercles; subcaudals transversely enlarged. Description of the holotype. Adult male, snout-vent length (SVL) 69.2 mm; body elongate (TrunkL/SVL 0.41); head elongate (HL/SVL 0.28), relatively narrow (HW/HL 0.63), depressed (HH/HL 0.33), distinct from neck; loreal region inflated, posterior nasal region concave; snout long (SE/HL 0.44), obtuse, longer than diameter of orbit (OD/SE 0.63); snout scales small, rounded, homogeneous, granular, larger than those on frontal and parietal regions; eye large (OD/HL 0.27), pupils vertical; eyelid fringe with tiny spines posteriorly; ear oval– shaped, small (EarL/HL 0.11); rostral wider than high with a medial suture; supranasals in contact medially; rostral bordered by first supralabial and nostril on each side; nares oval, surrounded by supranasal, rostral, first supralabial, and two enlarged postnasals; mental triangular, as wide as rostral (RW 3.0, MW 2.9); postmentals two, enlarged, in broad contact posteriorly, bordered by mental anteriorly, first two infralabials laterally; supralabials 10 / 11; infralabials 8 / 8. Dorsal scales small, granular to flattened; dorsal tubercles conical, round, present on occipital region, dorsum and tail base, those on the sides larger, each surrounded by 9 granular scales, in 20 irregular longitudinal rows at midbody; ventral scales smooth, medial scales 2 or 3 times larger than dorsal scales, round, subimbricate, in 32 longitudinal rows at midbody; lateral skin folds distinct, with tubercles; gular region with homogeneous, smooth scales; ventral scales between mental and cloacal slit 170; precloacal groove absent; a series of distinctly enlarged femoral scales present; femoral and precloacal pores 29, in a continuous row. \ Fore- and hind limbs moderately slender (ForeL/SVL 0.16, CrusL/SVL 0.20), femur longer than crus (FemurL/CrusL 1.34); dorsal surface of forelimbs with slightly developed tubercles; dorsal surface of hind limbs covered by distinctly developed tubercles; fingers and toes without distinct webbing; toe IV longer than finger IV (LD 4 P/LD 4 A 1.27); lamellae under fourth finger and fourth toe 18 / 18. Tail longer than snout–vent length (TaL 95.2 mm, TaL/SVL 1.38); postcloacal tubercles 5 / 5; dorsal surface of tail with distinct tubercles at base; subcaudals distinctly transversely enlarged, flat, smooth. Coloration in life. Ground color of dorsal surface of head and body greyish brown; dorsal head with irregular dark brown marking; nuchal loop dark brown, in U–shape, extending from posterior corner of eye, partly above tympanum to the neck; labials brown; five distinct dark transverse bands between limb insertions, normally with indentations in the posterior part of the dorsal bands at midbody; small tubercles at midbody brown; tubercles on sides yellow; dorsal surface of fore- and hind limbs with dark blotches; dorsal surface of tail grey with 15 dark bands, edged in white; posterior half of the tail white with only faint dark bands transversally; chin, throat, chest, belly and ventral surface of limbs greyish cream; ventral tail grey with dark and yellow blotches. Sexual dimorphism. The female differs from the males by the absence of femoral and precloacal pores (versus 29 pores in males) and the absence of hemipenial swellings at the tail base (see Tables 1 & 2). Comparisons. In the following we compare the new species with its congeners from Laos and neighbouring countries from the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on examination of specimens (see Appendix) and data obtained from the 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; Sumontha et al. 2015) (see Table 3). Morphologically, Cyrtodactylus soudthichaki sp. nov. closely resembles C. jaegeri Luu, Calame, Bonkowski, Nguyen & Ziegler and C. roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz in overall coloration and pattern. However, the new species can be distinguished from C. jaegeri by having fewer femoral and precloacal pores in males (29 versus 44 in C. jaegeri), more dorsal tubercle rows on body (19–20 versus 15–17 in C. jaegeri) and the absence of femoral and precloacal pores in the female (present in C. jaegeri), and from C. roesleri by having transverse dorsal bands between limb insertions with indentations in the posterior part of dorsal bands at the vertebral region (versus without indentations in C. roesleri) (see Ziegler et al. 2010), fewer ventral scale rows (32–33 versus 34–40 in C. roesleri), more femoral and precloacal pores in males (29 versus 20–28 in C. roesleri), and the absence of femoral pores in the female (present in C. roesleri). For further distinguishing characters see Table 4 and Fig. 4. TABLE 3. Morphological comparisons between Cyrtodactylus soudthichaki sp. nov. and its congeners from Laos and neighbouring countries in the Indochina region (compiled after Luu et al. 2014; Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014; Sumontha et al. 2015). Abbreviations are as follows: m = male; f= female; – = characters unobtainable from literature; * = tail regenerated, for other abbreviations see material and methods. . erythrops 78.4 83.0* 28 present present 10 + 9 + 9 – 16 20 blotched present (PP+FP) … …continued on the next page TABLE 3. (Continued) ……continued on the next page TABLE 3. (Continued) Cyrtodactylus soudthichaki sp. nov. has enlarged subcaudals, which are absent in the following species: C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. buchardi David, Teynié & Ohler, C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler, C. cucdongensis Schneider, Phung, Le, Nguyen & Ziegler, C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler, C. huynhi Ngo & Bauer, C. irregularis (Smith), C. phuocbinhensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, C. pseudoquadrivirgatus Rösler, Vu, Nguyen, Ngo & Ziegler, C. quadrivirgatus Taylor, C. ranongensis Sumontha, Pauwels, Panitvong, Kunya & Grismer, C. taynguyenensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, C. thuongae Phung, van Schingen, Ziegler & Nguyen, C. vilaphongi, and C. ziegleri Nazarov, Orlov, Nguyen & Ho. The new species has femoral and precloacal pores in males and thus differs from the following species which do not possess such pores in males: C. angularis (Smith), C. badenensis Nguyen, Orlov & Darevsky, C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, C. cucphuongensis Ngo & Chan, C. eisenmanae Ngo, C. grismeri Ngo, C. martini Ngo, C. nigriocularis Nguyen, Orlov & Darevsky, C. oldhami (Theobald), C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler, C. paradoxus (Darevsky & Szczerbak), C. puhuensis Nguyen, Yang, Le, Nguyen, Orlov, Hoang, Nguyen, Jin, Rao, Hoang, Che, Murphy & Zhang, C. saiyok Panitvong, Sumontha, Tunprasert & Pauwels, C. samroiyot Pauwels & Sumontha, C. sanook Pauwels, Sumontha, Latinne & Grismer, C. spelaeus, C. sumonthai Bauer, Pauwels & Chanhome, C. teyniei, C. wayakonei, C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome. The new species differs from C. astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels by its smaller size (SVL reaching 70.0 mm versus 108.3 mm), and fewer femoral and precloacal pores in males (29 versus 31–38); from C. auribalteatus Sumontha, Panitvong & Deein by having fewer ventral scale rows (32–33 versus 38–40), and more femoral and precloacal pores in males (29 versus 10–11); from C. bichnganae Ngo & Grismer by its smaller size (SVL 69.2 –70.0 mm versus 95.3–99.9 mm), and the absence of femoral and precloacal pores in the female (versus present); from C. brevipalmatus (Smith) by having fewer ventral scale rows (32–33 versus 35–44), more femoral and precloacal pores in males (29 versus 22), and the absence of femoral and precloacal pores in females (versus present); from C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen by its smaller size (SVL 69.2 –70.0 mm versus 90.4 –94.0 mm), having fewer ventral scale rows (32–33 versus 38–44), and more femoral and precloacal pores in males (29 versus 15); from C. chanhomeae Bauer, Sumontha & Pauwels by having fewer ventral scale rows (32–33 versus 36–38), fewer femoral and precloacal pores in males (29 versus 32), and the absence of femoral and precloacal pores in the female (versus present); from C. darevskii by its smaller size (SVL 69.2 –70.0 mm versus 84.6 –100.0 mm), having fewer ventral scale rows (32–33 versus 38–46), and fewer femoral and precloacal pores in males (29 versus 38–44); from C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya by its smaller size (SVL 69.2 –70.0 mm versus 76.2–84.2 mm), having fewer ventral scale rows (32–33 versus 40), and femoral and precloacal pores in a continuous row in males (versus discontinuous row); from C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya by its smaller size (SVL 69.2 –70.0 mm versus 78.4 mm), having more ventral scale rows (32–33 versus 28), and femoral and precloacal pores in a continuous row in males (versus discontinuous row); from C. huongsonensis Luu, Nguyen, Do & Ziegler by having fewer ventral scale rows (21–23 versus 41–48), more femoral and precloacal pores in males (29 versus 21–23), and the absence of femoral and precloacal pores in the female (versus present); from C. interdigitalis Ulber by its smaller size (SVL reaching 70.0 mm versus 80.0 mm), having fewer ventral scale rows (32–33 versus 37–42), and a slightly lower number of femoral and precloacal pores in males (29 versus 30–32); from C. intermedius (Smith) by its smaller size (SVL reaching 70.0 mm versus 85.0 mm) and by having fewer ventral scale rows (32–33 versus 40–50); from C. jarujini Ulber by its smaller size (SVL reaching 70.0 mm versus 90.0 mm) and having fewer femoral and precloacal pores in males (29 versus 52–54); from C. khammouanensis by having fewer ventral scales (32–33 versus 32–38) and fewer femoral and precloacal pores in males (29 versus 40–44); from C. khelangensis Pauwels, Sumontha, Panitvong & Varaguttanonda by its smaller size (SVL 69.2 –70.0 mm versus 72.8–95.3 mm in C. khelangensis), femoral and precloacal pores in a continuous row in males (versus discontinuous row), and the absence of femoral and precloacal pores in the female (versus present); from C. kingsadai Ziegler, Phung, Le & Nguyen by its smaller size (SVL reaching 70.0 mm versus 94.0 mm), having fewer ventral scales (32–33 versus 39–46), and more femoral and precloacal pores in males (29 versus 0–16); from C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels and C. lomyenensis in having fewer femoral and precloacal pores in males (29 versus 30–36 in C. lekaguli and 39–40 in C. lomyenensis), and the absence of femoral and precloacal pores in the female (versus present in C. lekaguli and C. lomyenensis); from C. multiporus by its smaller size (SVL reaching 70.0 mm versus 98.0 mm), having fewer ventral scale rows (32–33 versus 39–43), and fewer femoral and precloacal pores in males (29 versus 58–60); from C. phongnhakebangensis Ziegler, Rösler, Herrmann & Vu by its smaller size (SVL reaching 70.0 mm versus 96.3 mm), having fewer femoral and precloacal pores in males (29 versus 32–42), and the absence of femoral and precloacal pores in the female (versus present); from C. takouensis Ngo & Bauer by its smaller size (SVL 69.2 –70.0 mm versus 74.7–81.1 mm), having fewer ventral scales (32–33 versus 39–40) and more femoral and precloacal pores in males (29 versus 3–6); from C. tigroides Bauer, Sumontha & Pauwels by having a smaller size (SVL 69.2 –70.0 mm versus 74.3–83.2 mm) and more femoral and precloacal pores in males (29 versus 21); and from C. yangbayensis Ngo & Chan by having fewer ventral scale rows (32–33 versus 39–46) and more femoral and precloacal pores in males (29 versus 4–14). Distribution. Cyrtodactylus soudthichaki sp. nov. is currently known only from the type locality in Khun Don Mountain, Phou Hin Poun NPA, Khammouane Province, central Laos (Fig. 5). Etymology. This species is named in honour of Mr. Sisomphone Soudthichak, from the Natural Resources and Environment Department of Khammouane Province, Laos, who provided great support for our field research in Laos since 2013. As common names, we suggest Soudthichak’s Bent-toed Gecko (English) and Soudthichak’s Ki Chiem (Laotian). Natural history. The specimens were found between 19:00 and 21:00, on the branches of shrubs and karst boulders in a karst forest, approximately 0.3 m above the forest floor, between 150 and 170 m a.s.l. The karst forest included species of the dominated families Ebenaceae, Dracaenaceae, Arecaeae, Poaceae, Meliaceae, and Moraceae. The humidity at the time of collection was approximately 85 % and the air temperature ranged from 23 to 26 o C.

Cyrtodactylus soudthichaki sp. nov. (Figs. 1���3) Holotype. VFU R. 2015.18, adult male, Khun Don region (17 o 33.731 ���N, 104 o 52.360 ���E, elevation 167 m a.s.l.) within Phou Hin Poun NPA, Khammouane Province, central Laos, collected during night time between 19:00 and 21:00 on 4 April 2015 by V. Q. Luu and T. Calame. Paratypes. IEBR A. 2015.34, adult female and NUOL R- 2015.5, subadult male, bearing the same data as the holotype. Diagnosis. The new species can be distinguished from all other members of the genus Cyrtodactylus from the mainland Indochina region by a combination of the following characters: adult SVL 69.2 ���70.0 mm; dorsal head and neck with dark blotches; nuchal loop present; dorsum with five brown bands between limb insertions; 19 or 20 irregular rows of dorsal tubercles; 32 or 33 ventral scale rows; ventrolateral folds present with distinct tubercles; dorsal surface of hind limbs with tubercles; 29 precloacal and femoral pores in a continuous row in males, precloacal pores absent in the female; enlarged femoral and precloacal scales present; 4 or 5 postcloacal tubercles; subcaudals transversely enlarged. Description of the holotype. Adult male, snout-vent length (SVL) 69.2 mm; body elongate (TrunkL/SVL 0.41); head elongate (HL/SVL 0.28), relatively narrow (HW/HL 0.63), depressed (HH/HL 0.33), distinct from neck; loreal region inflated, posterior nasal region concave; snout long (SE/HL 0.44), obtuse, longer than diameter of orbit (OD/SE 0.63); snout scales small, rounded, homogeneous, granular, larger than those on frontal and parietal regions; eye large (OD/HL 0.27), pupils vertical; eyelid fringe with tiny spines posteriorly; ear oval��� shaped, small (EarL/HL 0.11); rostral wider than high with a medial suture; supranasals in contact medially; rostral bordered by first supralabial and nostril on each side; nares oval, surrounded by supranasal, rostral, first supralabial, and two enlarged postnasals; mental triangular, as wide as rostral (RW 3.0, MW 2.9); postmentals two, enlarged, in broad contact posteriorly, bordered by mental anteriorly, first two infralabials laterally; supralabials 10 / 11; infralabials 8 / 8. Dorsal scales small, granular to flattened; dorsal tubercles conical, round, present on occipital region, dorsum and tail base, those on the sides larger, each surrounded by 9 granular scales, in 20 irregular longitudinal rows at midbody; ventral scales smooth, medial scales 2 or 3 times larger than dorsal scales, round, subimbricate, in 32 longitudinal rows at midbody; lateral skin folds distinct, with tubercles; gular region with homogeneous, smooth scales; ventral scales between mental and cloacal slit 170; precloacal groove absent; a series of distinctly enlarged femoral scales present; femoral and precloacal pores 29, in a continuous row. \ Fore- and hind limbs moderately slender (ForeL/SVL 0.16, CrusL/SVL 0.20), femur longer than crus (FemurL/CrusL 1.34); dorsal surface of forelimbs with slightly developed tubercles; dorsal surface of hind limbs covered by distinctly developed tubercles; fingers and toes without distinct webbing; toe IV longer than finger IV (LD 4 P/LD 4 A 1.27); lamellae under fourth finger and fourth toe 18 / 18. Tail longer than snout���vent length (TaL 95.2 mm, TaL/SVL 1.38); postcloacal tubercles 5 / 5; dorsal surface of tail with distinct tubercles at base; subcaudals distinctly transversely enlarged, flat, smooth. Coloration in life. Ground color of dorsal surface of head and body greyish brown; dorsal head with irregular dark brown marking; nuchal loop dark brown, in U���shape, extending from posterior corner of eye, partly above tympanum to the neck; labials brown; five distinct dark transverse bands between limb insertions, normally with indentations in the posterior part of the dorsal bands at midbody; small tubercles at midbody brown; tubercles on sides yellow; dorsal surface of fore- and hind limbs with dark blotches; dorsal surface of tail grey with 15 dark bands, edged in white; posterior half of the tail white with only faint dark bands transversally; chin, throat, chest, belly and ventral surface of limbs greyish cream; ventral tail grey with dark and yellow blotches. Sexual dimorphism. The female differs from the males by the absence of femoral and precloacal pores (versus 29 pores in males) and the absence of hemipenial swellings at the tail base (see Tables 1 & 2). Comparisons. In the following we compare the new species with its congeners from Laos and neighbouring countries from the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on examination of specimens (see Appendix) and data obtained from the 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; Sumontha et al. 2015) (see Table 3). Morphologically, Cyrtodactylus soudthichaki sp. nov. closely resembles C. jaegeri Luu, Calame, Bonkowski, Nguyen & Ziegler and C. roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz in overall coloration and pattern. However, the new species can be distinguished from C. jaegeri by having fewer femoral and precloacal pores in males (29 versus 44 in C. jaegeri), more dorsal tubercle rows on body (19���20 versus 15���17 in C. jaegeri) and the absence of femoral and precloacal pores in the female (present in C. jaegeri), and from C. roesleri by having transverse dorsal bands between limb insertions with indentations in the posterior part of dorsal bands at the vertebral region (versus without indentations in C. roesleri) (see Ziegler et al. 2010), fewer ventral scale rows (32���33 versus 34���40 in C. roesleri), more femoral and precloacal pores in males (29 versus 20���28 in C. roesleri), and the absence of femoral pores in the female (present in C. roesleri). For further distinguishing characters see Table 4 and Fig. 4. TABLE 3. Morphological comparisons between Cyrtodactylus soudthichaki sp. nov. and its congeners from Laos and neighbouring countries in the Indochina region (compiled after Luu et al. 2014; Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014; Sumontha et al. 2015). Abbreviations are as follows: m = male; f= female; ��� = characters unobtainable from literature; * = tail regenerated, for other abbreviations see material and methods. . erythrops 78.4 83.0* 28 present present 10 + 9 + 9 ��� 16 20 blotched present (PP+FP) ��� ���continued on the next page TABLE 3. (Continued) ������continued on the next page TABLE 3. (Continued) Cyrtodactylus soudthichaki sp. nov. has enlarged subcaudals, which are absent in the following species: C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. buchardi David, Teyni�� & Ohler, C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. cattienensis Geissler, Nazarov, Orlov, B��hme, Phung, Nguyen & Ziegler, C. cucdongensis Schneider, Phung, Le, Nguyen & Ziegler, C. cryptus Heidrich, R��sler, Vu, B��hme & Ziegler, C. huynhi Ngo & Bauer, C. irregularis (Smith), C. phuocbinhensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, C. pseudoquadrivirgatus R��sler, Vu, Nguyen, Ngo & Ziegler, C. quadrivirgatus Taylor, C. ranongensis Sumontha, Pauwels, Panitvong, Kunya & Grismer, C. taynguyenensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, C. thuongae Phung, van Schingen, Ziegler & Nguyen, C. vilaphongi, and C. ziegleri Nazarov, Orlov, Nguyen & Ho. The new species has femoral and precloacal pores in males and thus differs from the following species which do not possess such pores in males: C. angularis (Smith), C. badenensis Nguyen, Orlov & Darevsky, C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, C. cucphuongensis Ngo & Chan, C. eisenmanae Ngo, C. grismeri Ngo, C. martini Ngo, C. nigriocularis Nguyen, Orlov & Darevsky, C. oldhami (Theobald), C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler, C. paradoxus (Darevsky & Szczerbak), C. puhuensis Nguyen, Yang, Le, Nguyen, Orlov, Hoang, Nguyen, Jin, Rao, Hoang, Che, Murphy & Zhang, C. saiyok Panitvong, Sumontha, Tunprasert & Pauwels, C. samroiyot Pauwels & Sumontha, C. sanook Pauwels, Sumontha, Latinne & Grismer, C. spelaeus, C. sumonthai Bauer, Pauwels & Chanhome, C. teyniei, C. wayakonei, C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome. The new species differs from C. astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels by its smaller size (SVL reaching 70.0 mm versus 108.3 mm), and fewer femoral and precloacal pores in males (29 versus 31���38); from C. auribalteatus Sumontha, Panitvong & Deein by having fewer ventral scale rows (32���33 versus 38���40), and more femoral and precloacal pores in males (29 versus 10���11); from C. bichnganae Ngo & Grismer by its smaller size (SVL 69.2 ���70.0 mm versus 95.3���99.9 mm), and the absence of femoral and precloacal pores in the female (versus present); from C. brevipalmatus (Smith) by having fewer ventral scale rows (32���33 versus 35���44), more femoral and precloacal pores in males (29 versus 22), and the absence of femoral and precloacal pores in females (versus present); from C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen by its smaller size (SVL 69.2 ���70.0 mm versus 90.4 ���94.0 mm), having fewer ventral scale rows (32���33 versus 38���44), and more femoral and precloacal pores in males (29 versus 15); from C. chanhomeae Bauer, Sumontha & Pauwels by having fewer ventral scale rows (32���33 versus 36���38), fewer femoral and precloacal pores in males (29 versus 32), and the absence of femoral and precloacal pores in the female (versus present); from C. darevskii by its smaller size (SVL 69.2 ���70.0 mm versus 84.6 ���100.0 mm), having fewer ventral scale rows (32���33 versus 38���46), and fewer femoral and precloacal pores in males (29 versus 38���44); from C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya by its smaller size (SVL 69.2 ���70.0 mm versus 76.2���84.2 mm), having fewer ventral scale rows (32���33 versus 40), and femoral and precloacal pores in a continuous row in males (versus discontinuous row); from C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya by its smaller size (SVL 69.2 ���70.0 mm versus 78.4 mm), having more ventral scale rows (32���33 versus 28), and femoral and precloacal pores in a continuous row in males (versus discontinuous row); from C. huongsonensis Luu, Nguyen, Do & Ziegler by having fewer ventral scale rows (21���23 versus 41���48), more femoral and precloacal pores in males (29 versus 21���23), and the absence of femoral and precloacal pores in the female (versus present); from C. interdigitalis Ulber by its smaller size (SVL reaching 70.0 mm versus 80.0 mm), having fewer ventral scale rows (32���33 versus 37���42), and a slightly lower number of femoral and precloacal pores in males (29 versus 30���32); from C. intermedius (Smith) by its smaller size (SVL reaching 70.0 mm versus 85.0 mm) and by having fewer ventral scale rows (32���33 versus 40���50); from C. jarujini Ulber by its smaller size (SVL reaching 70.0 mm versus 90.0 mm) and having fewer femoral and precloacal pores in males (29 versus 52���54); from C. khammouanensis by having fewer ventral scales (32���33 versus 32���38) and fewer femoral and precloacal pores in males (29 versus 40���44); from C. khelangensis Pauwels, Sumontha, Panitvong & Varaguttanonda by its smaller size (SVL 69.2 ���70.0 mm versus 72.8���95.3 mm in C. khelangensis), femoral and precloacal pores in a continuous row in males (versus discontinuous row), and the absence of femoral and precloacal pores in the female (versus present); from C. kingsadai Ziegler, Phung, Le & Nguyen by its smaller size (SVL reaching 70.0 mm versus 94.0 mm), having fewer ventral scales (32���33 versus 39���46), and more femoral and precloacal pores in males (29 versus 0���16); from C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels and C. lomyenensis in having fewer femoral and precloacal pores in males (29 versus 30���36 in C. lekaguli and 39���40 in C. lomyenensis), and the absence of femoral and precloacal pores in the female (versus present in C. lekaguli and C. lomyenensis); from C. multiporus by its smaller size (SVL reaching 70.0 mm versus 98.0 mm), having fewer ventral scale rows (32���33 versus 39���43), and fewer femoral and precloacal pores in males (29 versus 58���60); from C. phongnhakebangensis Ziegler, R��sler, Herrmann & Vu by its smaller size (SVL reaching 70.0 mm versus 96.3 mm), having fewer femoral and precloacal pores in males (29 versus 32���42), and the absence of femoral and precloacal pores in the female (versus present); from C. takouensis Ngo & Bauer by its smaller size (SVL 69.2 ���70.0 mm versus 74.7���81.1 mm), having fewer ventral scales (32���33 versus 39���40) and more femoral and precloacal pores in males (29 versus 3���6); from C. tigroides Bauer, Sumontha & Pauwels by having a smaller size (SVL 69.2 ���70.0 mm versus 74.3���83.2 mm) and more femoral and precloacal pores in males (29 versus 21); and from C. yangbayensis Ngo & Chan by having fewer ventral scale rows (32���33 versus 39���46) and more femoral and precloacal pores in males (29 versus 4���14). Distribution. Cyrtodactylus soudthichaki sp. nov. is currently known only from the type locality in Khun Don Mountain, Phou Hin Poun NPA, Khammouane Province, central Laos (Fig. 5). Etymology. This species is named in honour of Mr. Sisomphone Soudthichak, from the Natural Resources and Environment Department of Khammouane Province, Laos, who provided great support for our field research in Laos since 2013. As common names, we suggest Soudthichak���s Bent-toed Gecko (English) and Soudthichak���s Ki Chiem (Laotian). Natural history. The specimens were found between 19:00 and 21:00, on the branches of shrubs and karst boulders in a karst forest, approximately 0.3 m above the forest floor, between 150 and 170 m a.s.l. The karst forest included species of the dominated families Ebenaceae, Dracaenaceae, Arecaeae, Poaceae, Meliaceae, and Moraceae. The humidity at the time of collection was approximately 85 % and the air temperature ranged from 23 to 26 o C., Published as part of Luu, Vinh Quang, Calame, Thomas, Nguyen, Truong Quang, Bonkowski, Michael & Ziegler, Thomas, 2015, A new species of Cyrtodactylus (Squamata: Gekkonidae) from the limestone forest of Khammouane Province, central Laos, pp. 388-402 in Zootaxa 4058 (3) on pages 389-399, DOI: 10.11646/zootaxa.4058.3.6, http://zenodo.org/record/240248, {"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), 054 - 066. http: // dx. doi. org / 10.11646 / zootaxa. 3760.1.3","Nazarov, R., Poyarkov, N. A., Orlov, N. I., Nguyen, S. N., Milto, K. D., Martynov, A. A., Konstantinov, E. L. & Chulisov, A. S. (2014) A review of genus Cyrtodactylus (Reptilia: Sauria: Gekkonidae) in fauna of Laos with description of four new species. Toм 318, № 4, 2014, c, 391 - 423.","Nguyen, S. N., Yang, J. - X., Le, T. - N. T., Nguyen, L. T., Orlov, N. L., Hoang, C. V., Nguyen, T. Q., Jin, J. - Q., Rao, D. - Q., Hoang, T. N., Che, J., Murphy, R. W. & Zhang, Y. - P. (2014) DNA barcoding of Vietnamese bent-toed geckos (Squamata: Gekkonidae: Cyrtodactylus) and the description of a new species. Zootaxa, 3784 (1), 48 - 66. http: // dx. doi. org / 10.11646 / zootaxa. 3784.1.2","Schneider, N., Nguyen, T. Q., Le, M. D, Nophaseud, L., Bonkowski, M. & Ziegler, T. (2014) A new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of northern Laos. Zootaxa, 3835 (1), 80 - 96. http: // dx. doi. org / 10.11646 / zootaxa. 3835.1.4","Sumontha, M., Pauwels, O. S. G., Panitvong, N., Kunya, K. & Grismer, J. L. (2015) A new lowland forest Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Ranong Province, peninsular Thailand. Zootaxa, 3911 (1), 106 - 118. http: // dx. doi. org / 10.11646 / zootaxa. 3911.1.6","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, 20 - 36."]}

Cyrtodactylus vilaphongi sp. nov. (Figs. 2���4) Paratype. NUOL R- 2013.5, adult female, collected by Truong Quang Nguyen, Nicole Schneider, Liphone Nophaseud, Vilaphong Kanyasone on 10 August 2013, the same locality as the holotype. Diagnosis. The new species can be distinguished from all congeners on the basis of the following combination of characters: maximum SVL 86.1 mm, supralabials 9 or 10; infralabials 7���9; dorsal tubercles in 15 or 16 rows at midbody; ventral scale rows 34���36 at midbody; precloacal groove absent; femoral scales not distinctly enlarged; precloacal pores absent in females (unknown in males); subdigital lamellae under the fourth finger 18 or 19, under the fourth toe 18���20; subcaudals not transversally enlarged; dorsal bands white, 4 or 5 between limb insertions plus another one between hind limbs; tail banded. Description of the holotype. Adult female, SVL 86.1 mm, tail regenerated (TL 61.2 mm); rostral squareshaped, wider than high (RW 4.1 mm, RH 2.4 mm), medially with a straight, vertical suture, in contact with nasorostral, nare, and first supralabial on each side; supralabials 9 or 10; supralabials separated from orbit by 3 or 4 rows of scales; nares in contact with rostral, nasorostral, supranasal, two postnasals, and first supralabial; internasals absent; scales on snout distinctly larger than on upper head; loreal region slightly upraised; scales between fifth supralabials across the dorsal head surface in 30 rows; scales between anterior corners of eyes 32; interorbital region with small, round, convex scales, outer ones more oval; scales in postorbital region distinctly smaller (ca. half the size) than snout scales, round and convex, interrupted by a few small tubercles, in 3 or 4 rows laterally; dorsal surface of head without enlarged tubercles; pupil vertical; spinous ciliaria absent; ear opening vertical, oval; mental triangular, slightly narrower than rostral (MW: 3.6 mm), in contact with two postmentals and the first infralabial on each side; infralabials 8 or 9; postmentals surrounded by first infralabial on each side and six granular scales posteriorly, two outer ones enlarged; gular scales granular. Dorsal scales small; dorsal tubercles in 15 longitudinal rows at midbody, separated from each other by 2 or 3 dorsal scales and surrounded by 9���11 granular scales; tubercles on flanks slightly smaller than dorsal tubercles; ventrolateral folds slightly developed, with tubercles; ventral scales round, imbricated, 2 or 3 times larger than gular and throat scales, twice the size of dorsal scales; ventral scales 34, plus 122 dorsal scales around midbody; scales between mental and cloacal slit 165; dorsal surface of limbs with tubercles, distinctly higher counts on hind limbs; distinctly enlarged femoral scales absent; fingers and toes free of webbing; relative finger lengths IVariation. The paratype largely corresponds with the description of the holotype. For measurements, scalation, and color pattern variation see Fig. 2 and Table 3. Neck band is not interrupted in the paratype. The original tail has nine white dorsal bands plus a bright tip, dark brown ventrally, with dorsal tubercles at the base only, and subcaudals not transversally enlarged. Characters IEBR A. 2013.103 (female) NUOL R- 2013.5 (female) Measurements (mm) SVL 86.1 60.9 TL 61.2 * 68.1 AG 36.6 25.1 HL 23.6 16.6 HW 17.4 13.0 HH 9.7 6.9 SE 10.2 7.5 EE 7.4 4.9 IND 2.9 2.5 IOD 6.7 5.6 OD 5.5 4.1 ED 1.7 1.6 ForeaL 13.2 10.5 SL 14.9 12.4 LeF 4 7.5 5.5 LeT 4 9.6 6.8 RW 4.1 2.9 ......continued on the next page Distribution. Cyrtodactylus vilaphongi is currently known only from the type locality in Luang Prabang Province, Laos (Fig. 5). Etymology. The new species is named in honour of Mr. Vilaphong Kanyasone, a staff of Provincial Natural Resources and Environmental Office of Luang Prabang Province, Laos, who collected the holotype and provided a great support for our field research in Laos. Ecological notes. Cyrtodactylus vilaphongi inhabits disturbed secondary limestone forest near a residential area at elevations between 577 and 597 m a.s.l. Specimens were found at night, on karst cliff and rock boulder ca. 0.5 m above the ground (Fig. 6). Air temperature at the site was 24���30 o C and humidity ranged from 72 to 86 %., Published as part of Schneider, Nicole, Nguyen, Truong Quang, Le, Minh Duc, Nophaseud, Liphone, Bonkowski, Michael & Ziegler, Thomas, 2014, A new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of northern Laos in Zootaxa 3835 (1), DOI: 10.11646/zootaxa.3835.1.4, http://zenodo.org/record/224612

Luu, Vinh Quang, Calame, Thomas, Nguyen, Truong Quang, Le, Minh Duc, Bonkowski, Michael, Ziegler, Thomas (2014): A new species of the Gekko japonicus group (Squamata: Gekkonidae) from central Laos. Zootaxa 3895 (1): 73-88, DOI: http://dx.doi.org/10.11646/zootaxa.3895.1.4

Cyrtodactylus vilaphongi sp. nov. (Figs. 2–4) Paratype. NUOL R- 2013.5, adult female, collected by Truong Quang Nguyen, Nicole Schneider, Liphone Nophaseud, Vilaphong Kanyasone on 10 August 2013, the same locality as the holotype. Diagnosis. The new species can be distinguished from all congeners on the basis of the following combination of characters: maximum SVL 86.1 mm, supralabials 9 or 10; infralabials 7–9; dorsal tubercles in 15 or 16 rows at midbody; ventral scale rows 34–36 at midbody; precloacal groove absent; femoral scales not distinctly enlarged; precloacal pores absent in females (unknown in males); subdigital lamellae under the fourth finger 18 or 19, under the fourth toe 18–20; subcaudals not transversally enlarged; dorsal bands white, 4 or 5 between limb insertions plus another one between hind limbs; tail banded. Description of the holotype. Adult female, SVL 86.1 mm, tail regenerated (TL 61.2 mm); rostral squareshaped, wider than high (RW 4.1 mm, RH 2.4 mm), medially with a straight, vertical suture, in contact with nasorostral, nare, and first supralabial on each side; supralabials 9 or 10; supralabials separated from orbit by 3 or 4 rows of scales; nares in contact with rostral, nasorostral, supranasal, two postnasals, and first supralabial; internasals absent; scales on snout distinctly larger than on upper head; loreal region slightly upraised; scales between fifth supralabials across the dorsal head surface in 30 rows; scales between anterior corners of eyes 32; interorbital region with small, round, convex scales, outer ones more oval; scales in postorbital region distinctly smaller (ca. half the size) than snout scales, round and convex, interrupted by a few small tubercles, in 3 or 4 rows laterally; dorsal surface of head without enlarged tubercles; pupil vertical; spinous ciliaria absent; ear opening vertical, oval; mental triangular, slightly narrower than rostral (MW: 3.6 mm), in contact with two postmentals and the first infralabial on each side; infralabials 8 or 9; postmentals surrounded by first infralabial on each side and six granular scales posteriorly, two outer ones enlarged; gular scales granular. Dorsal scales small; dorsal tubercles in 15 longitudinal rows at midbody, separated from each other by 2 or 3 dorsal scales and surrounded by 9–11 granular scales; tubercles on flanks slightly smaller than dorsal tubercles; ventrolateral folds slightly developed, with tubercles; ventral scales round, imbricated, 2 or 3 times larger than gular and throat scales, twice the size of dorsal scales; ventral scales 34, plus 122 dorsal scales around midbody; scales between mental and cloacal slit 165; dorsal surface of limbs with tubercles, distinctly higher counts on hind limbs; distinctly enlarged femoral scales absent; fingers and toes free of webbing; relative finger lengths I Coloration in life: Dorsal head dark brown, marbled in light yellow; each side with a blackish brown stripe stretching from the posterior corner of the eye to the neck, connecting with the neck band on the right side, interrupted on the left side; irregular dark blotches with light margins present between eyes, on occiput and on postorbital region; ciliaria bright yellow; iris yellowish brown, pupil slit edged in orange; neck band blackish brown, widened posteriorly, edged in yellow anteriorly and posteriorly; ground color of dorsum blackish brown with five transverse yellowish white bands between limb insertions; flanks and upper limbs with yellowish white spots, in irregular rows; tail base dark brown with a white band, regenerated part uniformly dark brown; ventral surface light beige. Variation. The paratype largely corresponds with the description of the holotype. For measurements, scalation, and color pattern variation see Fig. 2 and Table 3. Neck band is not interrupted in the paratype. The original tail has nine white dorsal bands plus a bright tip, dark brown ventrally, with dorsal tubercles at the base only, and subcaudals not transversally enlarged. Characters IEBR A. 2013.103 (female) NUOL R- 2013.5 (female) Measurements (mm) SVL 86.1 60.9 TL 61.2 * 68.1 AG 36.6 25.1 HL 23.6 16.6 HW 17.4 13.0 HH 9.7 6.9 SE 10.2 7.5 EE 7.4 4.9 IND 2.9 2.5 IOD 6.7 5.6 OD 5.5 4.1 ED 1.7 1.6 ForeaL 13.2 10.5 SL 14.9 12.4 LeF 4 7.5 5.5 LeT 4 9.6 6.8 RW 4.1 2.9 ......continued on the next page Distribution. Cyrtodactylus vilaphongi is currently known only from the type locality in Luang Prabang Province, Laos (Fig. 5). Etymology. The new species is named in honour of Mr. Vilaphong Kanyasone, a staff of Provincial Natural Resources and Environmental Office of Luang Prabang Province, Laos, who collected the holotype and provided a great support for our field research in Laos. Ecological notes. Cyrtodactylus vilaphongi inhabits disturbed secondary limestone forest near a residential area at elevations between 577 and 597 m a.s.l. Specimens were found at night, on karst cliff and rock boulder ca. 0.5 m above the ground (Fig. 6). Air temperature at the site was 24–30 o C and humidity ranged from 72 to 86 %.

Hemiphyllodactylus kiziriani sp. nov. (Figs. 2, 3) Holotype. IEBR A. 2014.3, adult male, collected by Truong Quang Nguyen, Nicole Schneider, and Liphone Nophaseud on 10 August 2013, on limestone wall of a small cave (19 o 48.708 ���N, 102 o06.188���E, 637 m a.s.l. elevation), near Ban Xieng Muak Village, Luang Prabang District, Luang Prabang Province, Laos. Paratypes. Nine specimens from the same locality as the holotype: three adult males IEBR A. 2014.4, VNMN A. 2014.1 and ZFMK 95702 and two adult females NUOL R- 2014.1, VNMN A. 2014.2, collected on 10 August 2013; two females IEBR A. 2014.5, NUOL R- 2014.2, collected on 7 August (19 o 48.629 ���N, 102 o06.124���E, 594 a.s.l. m); ZFMK 95703, adult female, collected on 11 August (19 o 48.813 ���N, 102 o05.887���E, 628 m a.s.l. elevation); ZFMK 95704, adult female, collected on 16 August (19 o 48.772 ���N, 102 o06.181���E, 597 m a.s.l. elevation). Diagnosis. A bisexual taxon; SVL of adult males 35.1���40.1 mm, of adult females 36.3���40.8 mm; dorsal scale rows 18���27; ventral scale rows 11���15; chin scales bordering mental and first infralabial distinctly enlarged; digital lamellae formulae 3-4 - 4 - 4 (forefoot) and 4 - 4 / 5 - 4 / 5 - 4 (hindfoot); femoral pores 0���4, total precloacal pores 10���13 in males, 8���10 pitted precloacal scales in females; cloacal spurs present in both sexes, 1 or 2; dorsal trunk pattern of dark brown with two rows of irregular transverse bands; dark lateral head stripe distinct; upper zone of flank with a dark brown stripe; caecum and gonadal ducts unpigmented. Description of the holotype. Body dorsolaterally flattened, size small SVL 40.1mm, tail regenerated (TaL 19 + 11.8 mm), trunk length (TrunkL) 19.7mm, head slightly longer than wide (HeadL 7.4mm, HeadW 6.7mm), eye moderate (EyeD 2.7mm), ear opening oblique (EarD 0.8mm), nare-eye length (NarEye 3.2mm), snout-eye length (SnEye 4.2mm), internarial distance (SnW 1.7mm). Proportions: TrunkL/SVL 49.1 %, HeadL/SVL 18.5 %, HeadW/SVL 16.7 %, HeadW/HeadL 90.5 %, SnEye/HeadL 56.8 %, NarEye/HeadL 43.2 %, EyeD/HeadL 36.5 %, SnW/HeadL 23 %, EyeD/NarEye 84.4 %, SnW/HeadW 25.4 %. Scalation. Rostral very large, wider than high (rostral width 1.7mm, rostral height 0.9mm), with a shallow, but distinct suture; supralabials 11 / 11, enlarged from rostral to below eye, smaller in subocular rictus; nare in contact with rostral, first supralabial, supranasal, and two nasals posteriorly on each side; supranasals separated from each other by three small granular internasals; snout flat, covered by granular scales; pupil vertical; ear opening oblique, oval, approximately 29.6 % of the eye diameter, without bordering enlarged scales; mental triangular, wider than long; infralabials 10 / 10; three enlarged postmentals, outer ones distinctly larger than the middle scale, in contact with mental and first infralabials anteriorly; nine chin scales, all enlarged and of same size; dorsal scale rows 21���23 at midbody (contained within one eye diameter), enlarged tubercles absent, ventral scales larger than dorsal scales in 13 or 14 rows at midbody (contained within one eye diameter); dorsal surface of fore and hindlimbs covered with granular scales; terminal two phalanges free, claws absent on first finger, minute 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 - 4 - 4 (forefoot) and 4 - 4-5 - 4 (hindfoot); femoral and precloacal pore series continuous, 13 in total, asymmetrically arranged (femoral pores four on the left side, absent on the right side); cloacal spurs one; lamellae five on first fingers, five on first toes; dorsal caudal scales granular; subcaudals flat, slightly larger than dorsal caudal scales. Coloration in preservative. Ground color of dorsal surface of head and body yellowish grey; two dark brown streaks originating from posterior corner of eye on each side, the upper one extending to the neck but not forming a nuchal loop, the lower one connecting with a dark dorsolateral stripe extending along the flank to tail base; a dark brown band present on neck; neck and dorsum with a row of vertebral light spots; two rows of narrow dark brown bands present along dorsum but not connected with each other; dorsal surface of limbs grey with dark bars; upper and lower lips with dark bars; distinct postsacral mark with middorsal dark brown spot, bordered posteriorly and laterally by an U-shaped arm which can extend to posterior edge of abdomen; dorsal tail base with dark bands, generated part of the tail dark grey; throat, venter, and precloacal region cream, outer area with small dark brown dots; caecum and testis white, unpigmented. For color and pattern in life see Fig. 2. Variation. The scale counts vary among the type series: scales between supranasals two or three; supralabials 10 or 11; infralabials 9���11; chin scales 6���9; dorsal scale rows 18���27; ventral scale rows 11���15; cloacal spurs present in both sexes, 1 or 2, more distinct in males; femoral pores absent in all paratypes, precloacal pores 10���13 in males and 8���10 pitted precloacal scales in females; dorsal bands more distinct in two females (IEBR A. 2014.5, ZFMK 95704) (see Tables 3, 4). TABLE 2. Uncorrected (���p���) distance matrix showing percentage pairwise genetic divergence (ND 2) between Hemiphyllodactylus species. ���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� Species name 1 2 3 4 5 6 7 8 9 10 11 12 13 14 H. sp. nov. 2 (JN 393936) - H. sp. nov. 8 (JN 393949) 24.6 - H. sp. nov. 9 (JN 393935) 28.1 25.9 - H. aurantiacus (JN 393933) 24.8 24.5 20.4 - H. dushanensis (FJ 971017) 26.6 28.0 29.5 32.1 - H. ganoklonis (JN 393950) 19.1 25.8 28.2 25.6 30.0 - H. jinpingensis (FJ 971048) 27.3 18.7 28.3 24.6 29.6 28.3 - H. kiziriani sp. nov. 28.6-28.9 27.2-27.4 25.6-25.7 25.3-25.5 20.7 28.8-29.1 27.2-27.6 - (KJ 676800 - 2) H. longlingensi (FJ 971049) 26.8 20.2 28.0 26.9 26.4 28.6 18.4 26.1-26.4 - . H. titiwangsaensis (JN 393934) 28.4 27.6 28.6 29.3 30.0 28.2 29.0 30.7-31.2 28.1 - . H. titiwangsaensis (FJ 971050) 28.8 28.0 29.3 29.2 30.0 27.8 29.0 31.8-32.3 28.1 0.0 - . H. typus (GQ 257745) 19.3 26.2 29.0 25.2 30.3 17.7 28.0 27.6-27.7 29.3 29.6 29.8 - . H. yunnanensis (FJ 971044) 25.9 26.3 27.6 29.9 23.1 28.3 27.6 22.1-22.4 25.4 28.3 28.3 27.6 - . H. zugi (KF 575151 - 3) 26.4-26.6 25.8-26.2 27.0- 27.3 27.8-27.9 6.1-6.3 28.7-29.5 30.0- 30.1 20.1-20.6 26.1-26.7 29.7-30.2 30.1-30.5 29.2-29.3 22.8-22.9 - Table 4. Scalation of the type series of Hemiphyllodactylus kiziriani sp. nov. from Luang Prabang Province, Laos. Etymology. We name the new species in honour of Dr. David A. Kizirian, American Museum of Natural History (New York, USA), in recognition of his contribution to herpetological research in the Indochina region. As common names we suggest Kizirian���s Slender Gecko (English), Kizirians Halbblattfingergecko (German), and Th���ch s��ng d���p ki-zi-ri-an (Vietnamese). Natural history. Hemiphyllodactylus kiziriani inhabits disturbed secondary limestone forests near a residential area at elevations between 590 and 640 m. Specimens were found at night on tree bark and limestone cliffs near cave entrances or on a limestone boulder near a forest path, ca. 0.2���1.2 m above the ground. Two female paratypes (IEBR A. 2014.5, NUOL R- 2014.2) were gravid with two shelled eggs each. Distribution. The species is currently known only from Luang Prabang Province in northern Laos (Fig. 4)., Published as part of Nguyen, Truong Quang, Botov, Andreas, Le, Minh Duc, Nophaseud, Liphone, Zug, George, Bonkowski, Michael & Ziegler, Thomas, 2014, A new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Laos, pp. 45-56 in Zootaxa 3827 (1) on pages 48-55, DOI: 10.11646/zootaxa.3827.1.4, http://zenodo.org/record/286482

Cyrtodactylus jaegeri sp. nov. (Figs. 1���3) Holotype. IEBR A. 2013.55, adult male, collected on 29 April 2012 by Thomas Calame and Peter J��ger on a karst wall of a karstic massif, ca. 1.5 m above the forest floor, in a mixed secondary forest of hardwoods and shrubs near Thakhek Town (17 o 27.44 ���N, 104 o 55.44 ���E), Khammouane Province, Laos, at an elevation of 170 m a.s.l. Paratype. NUOL R��� 2013.1, adult female, the same collection data as the holotype. Diagnosis. A moderately sized Cyrtodactylus with a maximum SVL reaching 68.5 mm, distinguished from its congeners by a combination of the following characters: 1) dorsal pattern consisting of a dark nuchal loop and four narrow brown body bands between limb insertions; 3) dorsal tubercles in 15���17 irregular rows; 4) ventrals in 31��� 32 longitudinal rows at midbody; 5) lateral skin folds present with interspersed tubercles; 6) precloacal���femoral pores 44 in the male, in a continuous series; 7) enlarged femoral scales and precloacal scales present; 8) postcloacal tubercles 3���6; and 9) subcaudals transversely enlarged. Description of the holotype. Adult male, SVL 60 mm, body slender (TrunkL/SVL 0.41); head elongate (HL/ SVL 0.31), relatively narrow (HW/HL 0.61), depressed (HH/HL 0.4), distinguished from neck; loreal region inflated, frontonasal and posterior nasal regions concave; snout long (SE/HL 0.39), pointed, longer than diameter of orbit (OD/SE 0.65); snout scales small, rounded, homogeneous, granular, larger than those on frontal and parietal regions; eye large (OD/HL 0.26) with dark blue colour, pupils vertical; eyelid fringe with tiny spines posteriorly; ear in oval shape, small (EarL/HL 0.1); rostral wider than high (RW 2.9 mm, RH 1.3 mm, RW/RH 2.23) with a median suture; supranasals separated by a hexagonal internasal; rostral bordered by first supralabial and nostril on each side; nares oval, surrounded by supranasal, rostral, first supralabial, and two enlarged postnasals; mental triangular, slightly wider than rostral; postmentals in one pair, enlarged, longer than wide, in broad contact posteriorly, bordered by mental anteriorly, first two infralabials laterally, and one pair of distinctly enlarged gular scales posteriorly, which separated from each other by three small gular scales; supralabials 10 / 10; infralabials 9 / 9. Dorsal scales granular to flattened; dorsal tubercles round, conical, present on occipital region, back and tail base, surrounded by 9���10 granular scales, in 15 irregular longitudinal rows at midbody; ventral scales smooth, medial scales 2���3 times larger than dorsal scales, round, subimbricate, largest posteriorly, in 31 longitudinal rows at midbody; ventrolateral folds present with interspersed tubercles; gular region with homogeneous smooth scales; precloacal groove absent; a series of distinctly enlarged femoral scales present; femoral and precloacal pores 44, in a continuous series. Fore and hind limbs moderately slender (ForeL/SVL 0.17, CrusL/SVL 0.21); dorsal forelimbs with slightly developed tubercles; dorsal hindlimb covered by distinctly developed tubercles; fingers and toes without distinct webbing; lamellae under fourth finger 18 / 17, under fourth toe 19 / 19. Tail longer than snout���vent length (TaL 82.4 mm, TaL/SVL 1.37); postcloacal tubercles 5 / 6; dorsal tail bearing distinct tubercles at base; subcaudals distinctly transversely enlarged, flat, smooth. Coloration in ethanol. Ground coloration of dorsal head and back light brown; nuchal loop distinct, in U��� shape, from posterior corner of eye, running partly above tympanum to the neck, dark brown, edged in white posteriorly; body bands between limb insertions four and another one at the tail base, all thin, dark brown, edged in white posteriorly; dorsal tail brown with four dark brown bands in the anterior part, edged in white posteriorly, the bands indistinct in the posterior part; chin, throat, and belly cream; ventral tail dark brown; upper and lower lips with dark brown bars. For coloration in life see Fig. 2. Sexual dimorphism. The female paratype differs from the male holotype in having a larger size (SVL 68.5 mm vs. 60.0 mm), 21 femoral and precloacal-pitted scales in a continuous series, and the absence of an internasal scale as well as hemipenial swellings at the tail base (see Table 1). Comparisons. We compare the new species with its congeners from Laos and neighbouring countries in the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on examination of specimens (see Appendix) and data obtained from the literature (Table 2). Cyrtodactylus jaegeri sp. nov. has enlarged subcaudals, which are absent in C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, 2012, C. buchardi David, Teyni�� & Ohler, 2004, C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, 2012, C. cattienensis Geissler, Nazarov, Orlov, B��hme, Phung, Nguyen & Ziegler, 2009, C. cryptus Heidrich, R��sler, Vu, B��hme & Ziegler, 2007, C. huynhi Ngo & Bauer, 2008, C. irregularis (Smith, 1921), C. phuocbinhensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, 2013, C. pseudoquadrivirgatus R��sler, Vu, Nguyen, Ngo & Ziegler, 2008, C. quadrivirgatus Taylor, 1962, C. taynguyenensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, 2013, and C. ziegleri Nazarov, Orlov, Nguyen & Ho, 2008. Cyrtodactylus jaegeri sp. nov. differs from C. angularis (Smith, 1921) by having fewer ventral scale rows (31���32 versus 40���45 in C. angularis), the presence of femoral pores (absent in C. angularis), more femoral and precloacal pores in the male (44 vs. 3 in C. angularis). Cyrtodactylus jaegeri sp. nov. has 44 femoral and precloacal pores in the male and thus differs from Cyrtodactylus astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels, 2012 that has only 31���38 femoral and precloacal pores. Cyrtodactylus jaegeri sp. nov. differs from C. auribalteatus Sumontha, Panitvong & Deein, 2010 by having more enlarged femoral scales (18���19 vs. 5���7 in C. auribalteatus) and more femoral and precloacal pores in the male (44 vs. 10���11 in C. auribalteatus). Cyrtodactylus jaegeri sp. nov. can be distinguished from C. badenensis Nguyen, Orlov & Darevsky, 2006 by having enlarged femoral scales, the presence of femoral and precloacal pores (absent in C. badenensis), and more ventral scale rows (31���32 vs. 25���29 in C. badenensis). Cyrtodactylus jaegeri sp. nov. differs from C. bichnganae Ngo & Grismer, 2010 by having more enlarged femoral scales (18���19 vs. 11���13 in C. bichnganae) and more femoral and precloacal pores (44 vs. 28 in C. bichnganae) in males. Cyrtodactylus jaegeri sp. nov. has 44 femoral and precloacal pores in a continuous series and thus differs from C. brevipalmatus (Smith, 1923) and C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya, 2010, which are separated by poreless scales (6 + 9 + 7 in C. brevipalmatus and 6 + 5���6 + 6���7 in C. dumnuii). Cyrtodactylus jaegeri sp. nov. differs from C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen, 2007 by its smaller size (60.0��� 68.5 mm vs. 90.4 ���94.0 mm in C. caovansungi), fewer ventral scale rows (31���32 vs. 38���44 in C. caovansungi), and more femoral and precloacal pores in the male (44 vs. 15). Cyrtodactylus jaegeri sp. nov. differs from C. chanhomeae Bauer, Sumontha & Pauwels, 2003 in having fewer ventral scale rows (31���32 vs. 36��� 38) and more femoral and precloacal pores in the male (44 vs. 32). Cyrtodactylus jaegeri sp. nov. differs from C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, 2007 by its smaller size (SVL 60.0��� 68.5 mm vs. 90.9���99.3 mm), the presence of enlarged femoral scales and femoral pores (absent in C. chauquangensis). Cyrtodactylus jaegeri sp. nov. differs from C. cucphuongensis Ngo & Chan, 2011 by its smaller size (SVL 60���68.5 mm vs. 96 mm), fewer ventral scale rows (31���32 vs. 42), and the precence of femoral and precloacal pores (absent in C. cucphuongensis). Cyrtodactylus jaegeri sp. nov. has femoral and precloacal pores in males, which are absent in C. eisenmanae Ngo, 2008 and C. grismeri Ngo, 2008. Cyrtodactylus jaegeri sp. nov. differs from C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya, 2009 by having more ventral scale rows (31���32 vs. 28), 44 femoral and precloacal pores in a continuous series which are separated by poreless scales in C. erythrops (10 + 9 + 9). Cyrtodactylus jaegeri sp. nov. differs from C. huongsonensis Luu, Nguyen, Do & Ziegler, 2011 by having fewer ventral scale rows (31���32 vs. 41���48) and more femoral and precloacal pores in males (44 vs. 21���23). Cyrtodactylus jaegeri sp. nov. differs from C. interdigitalis Ulber, 1993 by having fewer ventral scale rows (31���32 vs. 37���42), more femoral and precloacal pores in males (44 vs. 30���32), and the presence of precloacal pitted scales in the female (absent in C. interdigitalis). Cyrtodactylus jaegeri sp. nov. differs from C. intermedius (Smith, 1917) by having fewer ventral scale rows (31���32 vs. 40���50) and more enlarged femoral scales (18���19 vs. 6���10). Cyrtodactylus jaegeri sp. nov. differs from C. jarujini Ulber, 1993 by its smaller size (60.0��� 68.5 vs. 85.0���90.0 in C. jarujini), fewer femoral and precloacal pores in males (44 vs. 52���54 in C. jarujini). Cyrtodactylus jaegeri sp. nov. differs from C. kingsadai Ziegler, Phung, Le & Nguyen, 2013 by its smaller size (60.0��� 68.5 vs. 83.0���94.0), fewer ventral scales (31���32 vs. 39���46), and more femoral and precloacal pores in males (44 vs. 0���16). Cyrtodactylus jaegeri sp. nov. differs from C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels, 2012 and C. lomyenensis Ngo & Pauwels, 2010 in having more femoral and precloacal pores in males (44 vs. 31���44 in C. lekaguli and 39���40 in C. lomyenensis), and fewer pitted scales in females (21 vs. 33���43 in C. lekaguli and 32 in C. lomyenensis). The new species differs from C. martini Ngo, 2011 by having fewer ventral scale rows (31���32 vs. 39���43 in C. martini) and the presence of femoral pores (absent in C. martini). Cyrtodactylus jaegeri sp. nov. differs from C. nigriocularis Nguyen, Orlov & Darevsky, 2006 in having smaller size (SVL 60���68.5 mm vs. 82.7���107.5 mm in C. nigriocularis), fewer ventral scale rows (31���32 vs. 42���49 in C. nigriocularis), and the presence of enlarged femoral scales and femoral pores (absent in C. nigriocularis). Cyrtodactylus jaegeri sp. nov. differs from C. oldhami (Theobald, 1876) by having more femoral and precloacal pores (44 vs. 1���4 in C. oldhami) and the dorsal color pattern (banded vs. striped and spotted body in C. oldhami). Cyrtodactylus jaegeri sp. nov. differs from C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler, 2011 and C. paradoxus (Darevsky & Szczerbak, 1997) by the presence of enlarged femoral scales and femoral pores (absent in C. pageli and C. paradoxus). Cyrtodactylus jaegeri sp. nov. differs from C. phongnhakebangensis Ziegler, R��sler, Herrmann & Vu, 2002 by its smaller size (SVL 60���68.5 mm vs. 85��� 96.3 mm in C. phongnhakebangensis) and a higher number of femoral and precloacal pores in males (44 vs. 32���42 in C. phongnhakebangensis). Cyrtodactylus jaegeri sp. nov. has 44 femoral and precloacal pores in the male and thus differs from C. roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz, 2010 (20���28). Cyrtodactylus jaegeri sp. nov. differs from C. sanook Pauwels, Sumontha, Latinne & Grismer, 2013 and C. sumonthai Bauer, Pauwels & Chanhome, 2002 by the presence of femoral pores (absent in C. sanook and C. sumonthai). Cyrtodactylus jaegeri sp. nov. differs from C. takouensis Ngo & Bauer, 2008 by its smaller size (SVL 60���68.5 mm vs. 74.7���81.1 mm in C. takouensis), fewer number of ventral scale rows (31���32 vs. 39���40), and more femoral and precloacal pores in males (44 vs. 3���6 in C. takouensis). Cyrtodactylus jaegeri sp. nov. differs from C. teyniei David, Nguyen, Schneider & Ziegler, 2011 by its smaller size (SVL 60���68.5 mm vs. 89.9 in C. teyniei), fewer number of ventral scale rows (31���32 vs. 38 in C. teyniei), and the presence of femoral pores (absent in C. teyniei). Cyrtodactylus jaegeri sp. nov. has femoral and precloacal pores which are lack in C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome 2004. Cyrtodactylus jaegeri sp. nov. has 44 femoral and precloacal pores in a contiuous series and thus differs from C. tigroides Bauer, Sumontha & Pauwels, 2003 which are separated by poreless scales (6 + 8 + 7 in males and 5 + 9 + 7 in females). Cyrtodactylus jaegeri sp. nov. has enlarged femoral scales and femoral pores which are absent in C. wayakonei Nguyen, Kingsada, R��sler, Auer & Ziegler, 2010. Cyrtodactylus jaegeri sp. nov. differs from C. yangbayensis Ngo & Chan, 2010 by its smaller size (SVL 60���68.5 mm vs. 78.5���92.3 mm in C. yangbayensis) and more femoral and precloacal pores in males (44 vs. 4���14 in C. yangbayensis). Distribution. Cyrtodactylus jaegeri sp. nov. is currently known only from the type locality in Khammouane Province, Laos (Fig. 4). Etymology. The new species is named in honour of our colleague Dr. Peter J��ger, arachnologist from the Research Institute Senckenberg, Frankfurt am Main, Germany, who collected the holotype. Ecological notes. The type specimens were found at night, on karst cliffs ca. 1.5 m above the ground in a small belt of the secondary vegetation in front of a limestone cliff, at an elevation of 170 m a.s.l., Published as part of Luu, Vinh Quang, Calame, Thomas, Bonkowski, Michael, Nguyen, Truong Quang & Ziegler, Thomas, 2014, A new species of Cyrtodactylus (Squamata: Gekkonidae) from Khammouane Province, Laos in Zootaxa 3760 (1), DOI: 10.11646/zootaxa.3760.1.3, http://zenodo.org/record/229973, {"references":["Nazarov, R., Poyarkov, N. A., Orlov, N. I., 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 - 24.","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, 621 - 627.","Geissler, P., Nazarov, R. A., Orlov, N. L., 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, 20 - 32.","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, 35 - 48.","Ngo, T. V. & Bauer, A. M. (2008) Descriptions of two new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) endemic to southern Vietnam. Zootaxa, 715, 27 - 42.","Smith, M. A. (1921) New or little - known Reptiles and Batrachians from Southern Annam (Indochina). Proceedings of Zoological Society of London, 1921, 423 - 440.","Nguyen, S. N., Le, N. T. T., Tran, D. T. A., Orlov, N. L., Lathrop, A., Macculloch, R. D., Le, D. T. T., Jin, J-Q., 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, 399 - 414. http: // dx. doi. org / 10.11646 / zootaxa. 3737.4.4","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.","Taylor, E. H. (1962) New oriental reptiles. University of Kansas Science Bulletin, 43, 209 - 263.","Nazarov, R. A., Orlov, N. L., Nguyen, S. N. & Ho, C. T. (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.","Grismer, L. L., Wood, P. L. Jr, Quah, E. S. H., Anuar, S., Muin, A., Sumontha, M., Ahmad, N., Bauer, A. M., Wangkulangkul, S., Grismer, J. L. & Pauwels, O. S. G. (2012) A phylogeny and taxonomy of the Thai - Malay Peninsula Bent - toed Geckos of the Cyrtodactylus pulchellus complex (Squamata: Gekkonidae): combined morphological and molecular analyses with descriptions of seven new species. Zootaxa, 3520, 1 - 55.","Sumontha, M., Panitvong, N. & Deein, G. (2010) Cyrtodactylus auribalteatus (Squamata: Gekkonidae), a new cave - dwelling gecko from Phitsanulok Province, Thailand. Zootaxa, 2370, 53 - 64.","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, 215 - 226.","Ngo, T. V. & Grismer, L. L. (2010) A new karst dwelling Cyrtodactylus (Squamata: Gekkonidae) from Son La Province, north - western Vietnam. Hamadryad, 35, 84 - 95.","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.","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, 41 - 50.","Orlov, N. L., Nguyen, T. Q., Nazarov, R. A., Ananjeva, 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]. Russian Journal of Herpetology, 14, 145 - 152.","Bauer, A. M., Sumontha, M. & Pauwels, O. S. G. (2003) Two new species of Cyrtodactylus (Reptilia: Squamata: Gekkonidae) from Thailand. Zootaxa, 376, 1 - 18.","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.","Ngo, T. V. & Chan, K. O. (2011) A new karstic cave - dwelling Cyrtodactylus Gray (Squamata: Gekkonidae) from northern Vietnam. Zootaxa, 3125, 51 - 63.","Bauer, A. M., Kunya, K., Sumontha, M., Niyomwan, P., Panitvong, N., Pauwels, O. S. G., Chanhome, L. & Kunya, T. (2009) Cyrtodactylus erythrops (Squamata: Gekkonidae), a new cave - dwelling gecko from Mae Hong Son Province, Thailand. Zootaxa, 2124, 51 - 62.","Luu, V. Q., Nguyen, T. Q., Do, H. Q. & Ziegler, T. (2011) A new Cyrtodactylus (Squamata: Gekkonidae) from Huong Son limestone forest, Hanoi, northern Vietnam. Zootaxa, 3129, 39 - 50.","Ulber, T. (1993) Bemerkungen uber cyrtodactyline Geckos aus Thailand nebst Beschreibungen von zwei neuen Arten (Reptilia: Gekkonidae). Mitteilungen aus dem Zoologischen Museum in Berlin, 69, 187 - 200. http: // dx. doi. org / 10.1002 / mmnz. 19930690202","Smith, M. A. (1917) Descriptions of new reptiles and a new batrachian from Siam. Journal of Natural History Society of Siam, 2, 221 - 225.","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, 432 - 466. http: // dx. doi. org / 10.11646 / zootaxa. 3686.4.2","Ngo, T. V. & Pauwels, O. S. G. (2010) A new cave - dwelling species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Khammouane Province, southern Laos. Zootaxa, 2730, 44 - 56.","Theobald, W. (1876) Descriptive Catalogue of the Reptiles of British India. Thacker, Spink & Co., Calcutta, xiii + 238 pp.","Schneider, N., Nguyen, T. Q., Schmitz, A., Kingsada, P., Auer, M. & Ziegler, T. (2011) A new species of karst dwelling Cyrtodactylus (Squamata: Gekkonidae) from northwestern Laos. Zootaxa, 2930, 1 - 21.","Darevsky, I. S. & Szczerbak, N. N. (1997) A new gecko of the genus Gonydactylus (Sauria: Gekkonidae) with a key to the species from Vietnam. Asiatic Herpetological Research, 7, 19 - 22.","Ziegler, T., Rosler, H., Herrmann, H. - W. & Vu, T. N. (2002) Cyrtodactylus phongnhakebangensis sp. n., ein neuer Bogenfingergecko aus dem annamitischen Karstwaldmassiv, Vietnam. Herpetofauna, 24, 11 - 25.","Ziegler, T., Nazarov, R., Orlov, N., Nguyen, T. Q., Vu, T. N., Dang, K. N., Dinh, T. H. & Schmitz, A. (2010) A third new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha - Ke Bang National Park, Vietnam. Zootaxa, 2413, 20 - 36.","Pauwels, O. S. G., Sumontha, M., Latinne, A. & Grismer, L. L. (2013) Cyrtodactylus sanook (Squamata: Gekkonidae), a new cave - dwelling gecko from Chumphon Province, southern Thailand. Zootaxa, 3635 (3), 275 - 285. http: // dx. doi. org / 10.11646 / zootaxa. 3635.3.7","Bauer, A. M., Pauwels, O. S. G. & Chanhome, L. (2002) A New Species of Cave - dwelling Cyrtodactylus (Squamata: Gekkonidae) from Thailand. Natural History Journal of Chulalongkorn University, 2 (2), 19 - 29.","David, P., Nguyen, T. Q., Schneider, N. & Ziegler, T. (2011) A new species of the genus Cyrtodactylus Gray, 1827 from central Laos (Squamata: Gekkonidae). Zootaxa, 2833, 29 - 40.","Pauwels, O. S. G., Bauer, A. M, Sumontha, M. & Chanhome, L. (2004) Cyrtodactylus thirakhupti (Squamata: Gekkonidae), a new cave - dwelling gecko from southern Thailand. Zootaxa, 772, 1 - 11.","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 - 16.","Ngo, T. V. & Chan, K. O. (2010) A new species of Cyrtodactylus Gray, 1826 (Squamata: Gekkonidae) from Khanh Hoa province, southern Vietnam. Zootaxa, 2504, 47 - 60."]}

Gekko thakhekensis sp. nov. (Figs. 2 and 4) Holotype. IEBR A. 2014.6, subadult male, collected on 29 April 2012 by Thomas Calame and Peter J��ger on a karst wall of a karstic massif, ca. 1.5 m above the forest floor, in a mixed secondary forest of hardwoods and shrubs near Thakhek Town (17 o 27.64 ���N, 104 o 55.24 ���E), Khammouane Province, Laos, at an elevation of 170 m a.s.l. Paratype. VFU R. 2014.9, adult male, collected on 0 2 June 2014 by Vinh Quang Luu and Thomas Calame on a liana near the wall of a karstic massif, ca. 2 m above the forest floor, in a mixed secondary forest of hardwoods and shrubs near Thakhek Town (17 o 27.707 ���N, 104 o 52.496 ���E), Khammouane Province, Laos, 4 km from the holotype locality at an elevation of 168 m a.s.l. Diagnosis. The new species differs from its relatives on the basis of the following combination of characters: size moderate (SVL 67.6���79.2 mm); nares in contact with rostral; internasals absent; postmentals enlarged; interorbital scales between anterior corners of the eyes 22���26; dorsal tubercles absent; ventral scales between mental and cloacal slit 165���174; midbody scale rows 110���116; ventral scale rows 32���40; subdigital lamellae on first toe 11���13, on fourth toe 14���15; finger and toe webbing present at base, about one fifth of length of digits; tubercles on upper surface of fore and hind limbs absent; precloacal pores 1���5 in males; postcloacal tubercles two; tubercles absent on dorsal surface of tail base; subcaudals enlarged; dorsal surface of body with greyish brown blotches. regenerated partially; for other abbreviations see material and methods). Description of holotype. Size moderate, SVL 67.6 mm, tail partially regenerated, TaL 66.7 mm, AG 29.3 mm; head longer than wide (HL 18.5 mm, HW 14.1 mm); rostral quadrangular, wider than high (RW 3.3 mm, RH 1.6 mm) and wider than mental (MW 2.2 mm), without suture; rostral in contact with first supralabial and supranasal; nostrils round, each surrounded by rostral, first supralabial, supranasal, and two enlarged nasals posteriorly; internasal absent; preorbitals 18; interorbitals 22; eye large (EE 4.5 mm, HL 18.5 mm), pupil vertical; ear opening oval, oblique, about 40 % of the eye diameter (maximum tympanum diameter 1.7 mm, horizontal eye diameter 4.5 mm; mental triangular, wider than long (MW 2.2 mm, ML 1.7 mm); postmentals two, relatively trapezoidal, twice longer than wide, and longer than length of mental, in contact with mental and first infralabials anteriorly, medial suture between; postmentals longer than the length of mental; postmental in contact with 5 gular scales posteriorly, outer gular scales larger than inner scales; supralabials 10 / 11; infralabials 10 / 9; dorsal scales on body smooth, round, granular and juxtaposed; lateral fold weakly developed; ventral scales much larger than dorsal scales, smooth, relatively hexagonal, imbricate, and largest in the middle of belly; ventrals between lateral folds 32; scales around midbody in 110 rows; ventral scales in a line between mental and cloacal slit 174; scales on upper and lower arm slightly enlarged; tubercles absent on dorsal surface of fore and hind limbs; scales on anterior and ventral parts of thigh larger than those on dorsal and posterior parts; enlarged femoral scales absent; fingers and toes basally webbed (about 1 / 5); subdigital lamellae under first finger 13 / 13, under fourth finger 15 / 15, under first toe 13 / 13, under fourth toe 15 / 15; precloacal pore one, precloacal scales enlarged; postcloacal tubercles 2 / 2, blunt; tail thickened at base, without tubercles on dorsal surface of tail base; dorsal caudal scales approximately twice the size than dorsal body scales, flat, in regular transverse rows; subcaudals flat, enlarged. Coloration in ethanol. Dorsal surface of head, body, limbs, and tail greyish brown with irregular vertebral blotches; nuchal surface with a light-colored patch, nuchal loop absent; upper eyelids greyish black; snout and interorbital region vermiculate; some small light spots present in temporal region and on sides of neck; neck with a light grey blotch; dorsum without vertebral stripe; some light and grey spots present on dorsal surface; a row of light spots present along lateral folds; limbs with small light spots and bars; throat, venter, and precloacal region yellowish cream with dark marbling; lower surface of tail brown. For coloration in life see Fig. 4. Variation. Measurements and scalation characters of the paratype are shown in Table 2. The following scale counts vary between the paratype and the holotype: interobitals 22���26, scale rows from mental to the front of cloacal slit 165���174, ventrals 32���40, and precloacal pores 1���5. Distribution. Gekko thakhekensis sp. nov. is currently known only from the type locality in Khammouane Province, Laos (Fig. 5). Ecological notes. The type specimens were collected at night, between 1.5 ���2.0 m above the ground in a small belt of the secondary vegetation in front of a limestone cliff, at elevations of 168���170 m a.s.l. Etymology. The specific epithet thakhekensis refers to the name of the type locality, Thakhek Town, Khammouane Province, Laos. Suggested common name: Thakhek Gecko. Character Gekko sp. nov. adleri auriverrucosus canhi chinensis japonicus hokouensis liboensis melli palmatus scabridus scientiadventura......continued on the next page TABLE 3. (Continued) ......continued on the next page TABLE 3. (continued) Character Gekko sp. nov. shibatai similignum subpalmatus swinhonis taibaiensis tawaensis vertebralis wenxianensis yakuensis. .....continued on the next page Comparisons. We compared the undescribed gecko species from Khammouane Province, central Laos with all other members of the Gekko japonicus group 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). Although only two male specimens were available for morphological comparisons, the undescribed species from Laos clearly differs from the remaining species of the G. japonicus group by a unique suite of features. The new Gekko species differs from the members of the G. japonicus group as follows: from G. adleri by lacking internasals (vs. one in G. adleri), dorsal tubercles absent (vs. present in G. adleri), fewer scale rows around midbody (110���116 vs. 123���144 in G. a d l e r i), fewer precloacal pores in males (1���5 vs. 17���21 in G. a d l e r i), more postcloacal tubercles (two vs. one in G. adleri), and tubercles on dorsal surface of hind limbs absent (present in G. adleri); from G. auriverrucosus Zhou & Liu by having a nostril touching rostral (not touching in G. auriverrucosus), postmentals enlarged (not enlarged in G. auriverrucosus), dorsal body tubercles absent (vs. present in G. auriverrucosus), and fewer precloacal pores in males (1���5 vs. 8���11 in G. auriverrucosus); from G. canhi R��sler, Nguyen, Doan, Ho & Ziegler by having fewer interorbitals (22���26 vs. 49���50 in G. canhi), internasal absent (vs. present in G. canhi), dorsal tubercles absent (vs. present in G. canhi), fewer scale rows around midbody (110���116 vs. 205���227 in G. canhi), fewer ventral scale rows (32���40 vs. 49���51 in G. c a n h i), and tubercles on dorsal surface of hind limbs absent (present in G. c a n h i); from G. chinensis Gray by lacking internasals (vs. one in G. chinensis), fewer interorbitals (22���26 vs. 35���48 in G. chinensis), dorsal tubercles absent (vs. present in G. chinensis), tubercles on dorsal surface of hind limbs absent (present in G. chinensis), fewer precloacal pores in males (1���5 vs. 17���27 in G. ch i n e n s i s), more postcloacal tubercles (two vs. one in G. chinensis), and tubercles on dorsal surface of tail absent (vs. present in G. chinensis); from G. japonicus (Schlegel) by having fewer interorbitals (22���26 vs. 32���35 in G. japonicus), postmentals enlarged (vs. not enlarged in G. japonicus), dorsal tubercles absent (vs. present in G. japonicus), fewer scale rows around midbody (110���116 vs. 130���144 in G. japonicus), tubercles on dorsal surface of fore and hind limbs absent (present in G. japonicus), fewer precloacal pores in males (1���5 vs. 6���9 in G. japonicus), and tubercles on dorsal surface of tail absent (vs. present in G. japonicus); from G. hokouensis Pope by lacking internasals (vs. one or two in G. hokouensis), fewer interorbitals (22���26 vs. 30���33 in G. hokouensis), postmentals enlarged (vs. not enlarged in G. hokouensis), dorsal tubercle rows absent (vs. present in G. hokouensis), fewer precloacal pores in males (1���5 vs. 5���9 in G. hokouensis), more postcloacal tubercles (two vs. one present in G. hokouensis), and tubercles on dorsal surface of tail absent (vs. present in G. hokouensis); from G. liboensis Zhao & Li by having fewer interorbitals (22���26 vs. 40 in G. l i b o e n s i s), dorsal tubercle rows absent (vs. present in G. liboensis), and more postcloacal tubercles (two vs. one in G. l i b o e n s i s); from G. melli Vogt by lacking internasals (vs. one present in G. melli), fewer interorbitals (22���26 vs. 34���40 in G. melli), postmentals enlarged (vs. not enlarged in G. melli), fewer scales in a line from mental to the front of cloacal slit (165���174 vs. 181���200 in G. melli), fewer scale rows around midbody (110���116 vs. 147���160 in G. melli), fewer ventral scale rows (32���40 vs. 43���49 in G. melli), fewer precloacal pores in males (1���5 vs. 9���11 in G. melli), and more postcloacal tubercles (two vs. one in G. melli); from G. palmatus by having dorsal tubercle rows absent (vs. present in G. palmatus), fewer precloacal pores in males (1���5 vs. 23���30 in G. palmatus), more postcloacal tubercles (two vs. one in G. palmatus), and tubercles on dorsal surface of tail absent (vs. present in G. palmatus); from G. scabridus Liu & Zhou by lacking internasals (vs. present in G. scabridus), fewer interorbitals (22���26 vs. 30 in G. scabridus), postmentals enlarged (vs. not enlarged in G. scabridus), dorsal tubercle rows absent (vs. present in G. scabridus), tubercles on dorsal surface of fore and hind limbs absent (present in G. scabridus), fewer precloacal pores in males (1���5 vs. 10���15 in G. scabridus), and tubercles on dorsal surface of tail absent (vs. present in G. scabridus); from G. scientiadventura by having fewer interorbitals (22���26 vs. 41���51 in G. scientiadventura), more scale rows from mental to cloacal slit (165���174 vs. 118���140 in G. scientiadventura), fewer scale rows around midbody (110���116 vs. 139���143 in G. scientiadventura), and fewer precloacal pores in males (1���5 vs. 5���8 in G. scientiadventura); from G. shibatai Toda, Sengoku, Hikida & Ota by having fewer interorbitals (22���26 vs. 37���52 in G. shibatai), postmentals enlarged (vs. not enlarged in G. shibatai), dorsal tubercle rows absent (vs. present in G. shibatai), more postcloacal tubercles (two vs. one in G. shibatai), and tubercles on dorsal surface of tail absent (vs. present in G. shibatai); from G. similignum Smith by having fewer interorbitals (22���26 vs. 46���48 in G. similignum), lacking internasals (vs. present in G. similignum), postmentals enlarged (vs. not enlarged in G. similignum), dorsal tubercle rows absent (vs. present in G. similignum), fewer scale rows around midbody (110���116 vs. 144���153 in G. similignum), fewer precloacal pores in males (1���5 vs. 17 in G. similignum), more postcloacal tubercles (two vs. one in G. similignum), and tubercles on dorsal surface of tail absent (vs. present in G. similignum); from G. subpalmatus G��nther by having fewer interorbitals (22���26 vs. 32 in G. subpalmatus), postmentals enlarged (vs. not enlarged in G. subpalmatus), internasals absent (vs. present in G. subpalmatus), fewer ventral scale rows (32���40 vs. 48 in G. subpalmatus), and fewer precloacal pores in males (1���5 vs. 5���11 in G. subpalmatus), and more postcloacal tubercles (two vs. one in G. subpalmatus); from G. swinhonis G��nther by having postmentals enlarged (vs. not enlarged in G. swinhonis), dorsal tubercle rows absent (vs. present in G. swinhonis), tubercles on dorsal surface of fore and hind limbs absent (present in G. swinhonis), and fewer precloacal pores in males (1���5 vs. 7���9 in G. swinhonis); from G. taibaiensis Song by having a larger size (SVL 79.2 vs. 69.0 mm in G. taibaiensis), more lamellae under first and fourth toes (11���13 vs. 6���7 and 14���15 vs. 7���8, respectively, in G. taibaiensis), and fewer precloacal pores in males (1���5 vs. 4���6 in G. taibaiensis); from G. tawaensis Okada by lacking internasals (vs. present in G. tawaensis), postmentals enlarged (vs. not enlarged in G. tawaensis), precloacal pores present (vs. absent in G. tawaensis), and more postcloacal tubercles (two vs. one in G. tawaensis); from G. vertebralis Toda, Sengoku, Hikida & Ota by having fewer interorbitals (22���26 vs. 35���50 in G. vertebralis), postmentals enlarged (vs. not enlarged in G. vertebralis), dorsal tubercle rows absent (vs. present in G. vertebralis), and tubercles on dorsal surface of tail absent (vs. present in G. vertebralis); from G. wenxianensis Zhou & Wang by lacking internasals (vs. present in G. wenxianensis), dorsal tubercle rows absent (vs. present in G. wenxianensis), fewer ventral scale rows (32���40 vs. 42���44 in G. wenxianensis), tubercles on dorsal surface of hind limbs absent (present in G. wenxianensis), and fewer precloacal pores in males (1���5 vs. 6���8 in G. wenxianensis); and from G. yakuensis Matsui & Okada by having internasals absent (vs. present in G. yakuensis), postmentals enlarged (vs. not enlarged in G. yakuensis), fewer precloacal pores in males (1���5 vs. 6���8 in G. yakuensis), more postcloacal tubercles (two vs. one in G. yakuensis), and tubercles on dorsal surface of tail absent (vs. present in G. yakuensis)., Published as part of Luu, Vinh Quang, Calame, Thomas, Nguyen, Truong Quang, Le, Minh Duc, Bonkowski, Michael & Ziegler, Thomas, 2014, A new species of the Gekko japonicus group (Squamata: Gekkonidae) from central Laos, pp. 73-88 in Zootaxa 3895 (1) on pages 76-86, DOI: 10.11646/zootaxa.3895.1.4, http://zenodo.org/record/287617, {"references":["Boulenger, G. A. (1907) Description of new lizard in the British Museum. Annals and Magazine of Natural History, London, 19, 486 - 489.","Ota, H., Lau, M. W., Weidenhofer, T., Yasukawa, Y. & Bogadek, A. (1995) Taxonomic review of the geckos allied to Gekko chinensis Gray 1842 (Gekkonidae, Reptilia) from China and Vietnam. Tropical Zoology, 8, 181 - 196. http: // dx. doi. org / 10.1080 / 03946975.1995.10539278","Rosler, H., Ziegler, T., Vu, N. T., Herrmann, H. W. & Bohme, W. (2005 \" 2004 \") A new lizard of the Genus Gekko Laurenti, 1768 (Squamata: Sauria: Gekkonidae) from the Phong Nha-Ke Bang National Park, Quang Binh Province, Vietnam. Bonner zoologische Beitrage, 53 (1 / 2), 135 - 148.","Rosler, H., Nguyen, Q. T., Doan, V. K., Ho, T. C., Nguyen, T. T. & Ziegler, T. (2010) A new species of the genus Gekko Laurenti (Squamata: Sauria: Gekkonidae) from Vietnam with remarks on G. japonicus (Schlegel). Zootaxa, 2329, 56 - 68.","Rosler, H., Bauer, A. M., Heinicke, M., Greenbaum, E., Jackman, T., Nguyen, Q. T. & Ziegler, T. (2011) Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae). Zootaxa, 2989, 1 - 50.","Yang, J. - H., Wang, Y. - Y., Zhang, T. - D., Sun, Y. - J. & Lin, S. - S. (2012) Genetic and morphological evidence on the species validity of Gekko melli Vogt, 1922 with notes on its diagnosis and range extension (Squamata: Gekkonidae). Zootaxa, 3505, 67 - 74.","Nguyen, Q. T., Wang, Y-Y., Yang, J-H., Lehmann, T., Le, M. D., Ziegler, T. & Bonkowski, M. (2013) A new species of the Gekko japonicus group (Squamata: Sauria: Gekkonidae) from the border region between China and Vietnam. Zootaxa, 3652 (5), 501 - 518. http: // dx. doi. org / 10.11646 / zootaxa. 3652.5.1"]}

Three rare nematode species of the family Belondiridae, originally described from and only known to occur in India are recorded for the first time in Vietnam: Axonchium thoubalicum, Belondira murtazai and Oxybelondira paraperplexa. It is the first report of these three genera in this country. The three species are described, including new morphological data, morphometrics and light microscope pictures. The male of O. paraperplexa is collected and described for the first time. It is characterized by its 1.54 mm long body, ad-cloacal pair of genital papillae situated at 9.0 µm from the cloacal aperture, only one ventromedian supplement located at 15 µm from the ad-cloacal pair within the range of spicules, spicules slightly curved ventrad and 42 µm long (7 times as long as wide and 2 times as long as cloacal body diameter), and tail 100 µm long (c = 15, c' = 5) and similar to that of the female.

Sphenomorphus sheai sp. nov. (Figs. 1, 2) Holotype. IEBR A. 2013.22 (Field number ASU 10715), adult female, collected on 4 June 2012 by K. V. Nguyen and R. W. Van Devender near Camp 1 (14 o 37.098 ���N, 108 o 30.177 ���E, elevation 1,025 m; datum WGS 84) from Kon Tum Plateau in the border area between Quang Ngai and Kon Tum provinces, southern Central Vietnam. Diagnosis. Size small (SVL 35.2 mm); prefrontals separated from each other; supraoculars four; enlarged nuchals in two pairs; lower eyelid scaly; supralabials six, first fused with nasal; primary temporals two; external ear openings absent; midbody scales in 20 rows; dorsal scales smooth, paravertebral scales 53, not widened; limbs short, pentadactyl; scales on dorsal surface of base of fourth toe in three rows; lamellae under fourth toe six; free margins of upper and lower eyelids not edged in white; dorsum and tail base bronze brown with dark dots; dorsolateral light stripe present on neck, shoulder and anterior part of dorsum; upper zone of lateral head and flank with a narrow black stripe. Description of holotype. Size small (SVL 35.2 mm), tail complete (TaL 58.5 mm), head longer than wide (HL 6.0 mm, HW 4.2 mm), SL 2.6 mm, SFlL 10.8 mm, AG 21.2 mm, limbs short (FlL 5.1 mm, HlL 7.5 mm); proportions: TaL/SVL 1.66, HL/SVL 0.17, FlL/SVL 0.14, HlL/SVL 0.21. Snout obtuse, rounded anteriorly; rostral wider than high, distinctly visible from above; supranasals absent; frontonasal wider than long, in contact with rostral, nasals, anterior loreals, and prefrontals; prefrontals separated from each other; frontal narrowing posteriorly, longer than the distance to the tip of snout, in contact with frontonasal, prefrontals, first and second supraoculars, and frontoparietals; frontoparietals in contact with each other anteriorly, and bordered by frontal, three posterior supraoculars, parietals, and interparietal; interparietal lozenge-shaped with a small transparent spot in posterior angle; parietals in contact posteriorly, posterolateral border surrounded by three scales on each side; enlarged nuchal scales in two pairs. Nostril in center of nasal; postnasals absent; loreals two, anterior larger than posterior; preocular single; presubocular single; supraciliaries seven, first largest, first to third in contact with first supraocular; supraoculars four, second widest, fourth supraocular followed by a small scale; postocular single; postsuboculars three, lower one in contact with fifth supralabial; primary temporals two, lower one in contact with sixth supralabial; secondary temporals two, upper one very large, in contact with parietal, overlapped by lower one; lower eyelid moveable, scaly, separated from supralabials by one row of small scales; supralabials six, anterior first fused with nasal, fifth and sixth enlarged, fourth below the eye; external ear openings absent. Mental wider than long, rounded anteriorly, in contact with anterior infralabial on each side and postmental; infralabials five; postmental undivided, in contact with mental, first infralabial and anterior pair of chinshields; three pairs of chinshields, anterior pair in contact with each other anteriorly, second pair separated from each other by a gular scale, and posterior pair separated from each other by three scales. Midbody scales in 20 rows; dorsal scales smooth, subequal to lateral and ventral scales, four scale rows between dark stripes on upper lateral zones; paravertebral scales 53, not widened; ventrals smooth, in 54 rows; precloacals four, inner scales overlapping outer ones, medial two enlarged, right scale overlapped by left scale; tail thick at base, median subcaudals widened. Limbs short, pentadactyl; second and fourth fingers equal in length, third longest; fore and hind limbs distinctly separated when adpressed along body wall; scales on dorsal surface of base of fourth toe in three rows, reducing to two at third or fourth scale and with terminal two scales single; subdigital lamellae smooth, numbering five under fourth finger and six under fourth toe Coloration in alcohol. Free margins of upper and lower eyelids not edged in white; dorsum and tail base bronze brown with fine dark dots; dorsal tail tip dark brown; upper lateral head and flank with narrow black stripe, one scale wide, running from nostril to anterior corner of eye and from behind the eye through hindlimb to tail base, interrupted by small light spots from posterior half of body; dorsolateral light stripe present on neck, shoulder and anterior part of dorsum; supralabials and infralabials with dark bars on sutures; arms and legs dark brown with light spots; chin and throat cream; venter and tail base cream; underside of tail tip with very small dark brown dots. Distribution. Sphenomorphus sheai sp. nov. is known only from the type locality in the border area between Quang Ngai and Kon Tumprovince, southern Central Vietnam (Fig. 3). Natural history. The holotype was collected about two hours after dark among leaf litter on a steep slope about three meters above a small stream surrounded by secondary forest at an elevation of 1025 m. The holotype contained four small white follicles in each ovary. Etymology. The new species is named in honour of Dr. Glenn Shea from the University of Sydney, Australia, in recognition of his outstanding contributions towards the systematics of scincid lizards. For the common names we suggest Shea���s Forest Skink (English), Th���n l���n ph��-n�� shea (Vietnamese), and Sheas Bodenskink (German). Comparisons. Based on specimen examination (Appendix) and data from the literature (e.g., Boulenger 1887; Smith 1935; Bourret 1939, 2009; Taylor 1963; Darevsky & Orlov 2005; Nguyen et al. 2011; Luu et al. 2013; Roy et al. 2013), we compare Sphenomorphus sheai with the following species of Sphenomorphus from China and mainland Southeast Asia which have midbody scale counts fewer than 26 rows: Sphenomorphus stellatus (Boulenger) (22���24 rows), S. tridigitus (Bourret) (18���20 rows), S. tetradactylus (Darevsky & Orlov) (20 rows). Sphenomorphus sheai can be distinguished from S. stellatus by having a smaller size (SVL 35.2 mm vs. 57���80 mm in S. stellatus), fewer lamellae under fourth toe (six vs. 18���23 in S. stellatus), fore and hind limbs distinctly separated when adpressed along body wall (vs. overlapped), and by the absence of dorsal large dark blotches (vs. presence). The new species differs from S. tridigitus and S. tetradactylus by having forelimbs pentydactyl (versus three and four in S. tridigitus and S. tetradactylus, respectively) and two primary temporals (vs. one in S. tridigitus and S. tetradactylus). Sphenomorphus sheai could be mistaken for several skink species of the genus Lygosoma, which also have short limbs and midbody scale rows fewer than 26, including L. anguinum (Theobald) (22), L. lineolatum (Stolizcka) (22), L. popae (Shreve) (24), L. punctata (Gmelin) (24���28), L. quadrupes (Linnaeus) (24���28), and L. veunsaiensis Geissler, Hartmann & Neang (22). However, Sphenomorphus sheai differs from all aforementioned species by the absence of supranasals (present in Lygosoma) and having the first supralabial fused with nasal (not fused in Lygosma) (see Geissler et al. 2012). Sphenomorphus sheai also resembles Scincella apraefrontalis Nguyen, Nguyen, B��hme & Ziegler by having a low midbody scale count, no ear openings, and first supralabial fused with nasal; however, the new species can be distinguished from latter by the presence of prefrontals and opaque window on lower eyelid which are absent in Scincella apraefrontalis (Nguyen et al. 2010)., Published as part of Nguyen, Truong Quang, Nguyen, Khoi Vu, Devender, Robert Wayne Van, Bonkowski, Michael & Ziegler, Thomas, 2013, A new species of Sphenomorphus Fitzinger, 1843 (Squamata: Sauria: Scincidae) from Vietnam in Zootaxa 3734 (1), DOI: 10.11646/zootaxa.3734.1.6, http://zenodo.org/record/223677

Nguyen, Truong Quang, Nguyen, Khoi Vu, Devender, Robert Wayne Van, Bonkowski, Michael, Ziegler, Thomas (2013): A new species of Sphenomorphus Fitzinger, 1843 (Squamata: Sauria: Scincidae) from Vietnam. Zootaxa 3734 (1), DOI: http://dx.doi.org/10.11646/zootaxa.3734.1.6

Sphenomorphus sheai sp. nov. (Figs. 1, 2) Holotype. IEBR A. 2013.22 (Field number ASU 10715), adult female, collected on 4 June 2012 by K. V. Nguyen and R. W. Van Devender near Camp 1 (14 o 37.098 ’N, 108 o 30.177 ’E, elevation 1,025 m; datum WGS 84) from Kon Tum Plateau in the border area between Quang Ngai and Kon Tum provinces, southern Central Vietnam. Diagnosis. Size small (SVL 35.2 mm); prefrontals separated from each other; supraoculars four; enlarged nuchals in two pairs; lower eyelid scaly; supralabials six, first fused with nasal; primary temporals two; external ear openings absent; midbody scales in 20 rows; dorsal scales smooth, paravertebral scales 53, not widened; limbs short, pentadactyl; scales on dorsal surface of base of fourth toe in three rows; lamellae under fourth toe six; free margins of upper and lower eyelids not edged in white; dorsum and tail base bronze brown with dark dots; dorsolateral light stripe present on neck, shoulder and anterior part of dorsum; upper zone of lateral head and flank with a narrow black stripe. Description of holotype. Size small (SVL 35.2 mm), tail complete (TaL 58.5 mm), head longer than wide (HL 6.0 mm, HW 4.2 mm), SL 2.6 mm, SFlL 10.8 mm, AG 21.2 mm, limbs short (FlL 5.1 mm, HlL 7.5 mm); proportions: TaL/SVL 1.66, HL/SVL 0.17, FlL/SVL 0.14, HlL/SVL 0.21. Snout obtuse, rounded anteriorly; rostral wider than high, distinctly visible from above; supranasals absent; frontonasal wider than long, in contact with rostral, nasals, anterior loreals, and prefrontals; prefrontals separated from each other; frontal narrowing posteriorly, longer than the distance to the tip of snout, in contact with frontonasal, prefrontals, first and second supraoculars, and frontoparietals; frontoparietals in contact with each other anteriorly, and bordered by frontal, three posterior supraoculars, parietals, and interparietal; interparietal lozenge-shaped with a small transparent spot in posterior angle; parietals in contact posteriorly, posterolateral border surrounded by three scales on each side; enlarged nuchal scales in two pairs. Nostril in center of nasal; postnasals absent; loreals two, anterior larger than posterior; preocular single; presubocular single; supraciliaries seven, first largest, first to third in contact with first supraocular; supraoculars four, second widest, fourth supraocular followed by a small scale; postocular single; postsuboculars three, lower one in contact with fifth supralabial; primary temporals two, lower one in contact with sixth supralabial; secondary temporals two, upper one very large, in contact with parietal, overlapped by lower one; lower eyelid moveable, scaly, separated from supralabials by one row of small scales; supralabials six, anterior first fused with nasal, fifth and sixth enlarged, fourth below the eye; external ear openings absent. Mental wider than long, rounded anteriorly, in contact with anterior infralabial on each side and postmental; infralabials five; postmental undivided, in contact with mental, first infralabial and anterior pair of chinshields; three pairs of chinshields, anterior pair in contact with each other anteriorly, second pair separated from each other by a gular scale, and posterior pair separated from each other by three scales. Midbody scales in 20 rows; dorsal scales smooth, subequal to lateral and ventral scales, four scale rows between dark stripes on upper lateral zones; paravertebral scales 53, not widened; ventrals smooth, in 54 rows; precloacals four, inner scales overlapping outer ones, medial two enlarged, right scale overlapped by left scale; tail thick at base, median subcaudals widened. Limbs short, pentadactyl; second and fourth fingers equal in length, third longest; fore and hind limbs distinctly separated when adpressed along body wall; scales on dorsal surface of base of fourth toe in three rows, reducing to two at third or fourth scale and with terminal two scales single; subdigital lamellae smooth, numbering five under fourth finger and six under fourth toe Coloration in alcohol. Free margins of upper and lower eyelids not edged in white; dorsum and tail base bronze brown with fine dark dots; dorsal tail tip dark brown; upper lateral head and flank with narrow black stripe, one scale wide, running from nostril to anterior corner of eye and from behind the eye through hindlimb to tail base, interrupted by small light spots from posterior half of body; dorsolateral light stripe present on neck, shoulder and anterior part of dorsum; supralabials and infralabials with dark bars on sutures; arms and legs dark brown with light spots; chin and throat cream; venter and tail base cream; underside of tail tip with very small dark brown dots. Distribution. Sphenomorphus sheai sp. nov. is known only from the type locality in the border area between Quang Ngai and Kon Tumprovince, southern Central Vietnam (Fig. 3). Natural history. The holotype was collected about two hours after dark among leaf litter on a steep slope about three meters above a small stream surrounded by secondary forest at an elevation of 1025 m. The holotype contained four small white follicles in each ovary. Etymology. The new species is named in honour of Dr. Glenn Shea from the University of Sydney, Australia, in recognition of his outstanding contributions towards the systematics of scincid lizards. For the common names we suggest Shea’s Forest Skink (English), Thằn lằn phê-nô shea (Vietnamese), and Sheas Bodenskink (German). Comparisons. Based on specimen examination (Appendix) and data from the literature (e.g., Boulenger 1887; Smith 1935; Bourret 1939, 2009; Taylor 1963; Darevsky & Orlov 2005; Nguyen et al. 2011; Luu et al. 2013; Roy et al. 2013), we compare Sphenomorphus sheai with the following species of Sphenomorphus from China and mainland Southeast Asia which have midbody scale counts fewer than 26 rows: Sphenomorphus stellatus (Boulenger) (22–24 rows), S. tridigitus (Bourret) (18–20 rows), S. tetradactylus (Darevsky & Orlov) (20 rows). Sphenomorphus sheai can be distinguished from S. stellatus by having a smaller size (SVL 35.2 mm vs. 57–80 mm in S. stellatus), fewer lamellae under fourth toe (six vs. 18–23 in S. stellatus), fore and hind limbs distinctly separated when adpressed along body wall (vs. overlapped), and by the absence of dorsal large dark blotches (vs. presence). The new species differs from S. tridigitus and S. tetradactylus by having forelimbs pentydactyl (versus three and four in S. tridigitus and S. tetradactylus, respectively) and two primary temporals (vs. one in S. tridigitus and S. tetradactylus). Sphenomorphus sheai could be mistaken for several skink species of the genus Lygosoma, which also have short limbs and midbody scale rows fewer than 26, including L. anguinum (Theobald) (22), L. lineolatum (Stolizcka) (22), L. popae (Shreve) (24), L. punctata (Gmelin) (24–28), L. quadrupes (Linnaeus) (24–28), and L. veunsaiensis Geissler, Hartmann & Neang (22). However, Sphenomorphus sheai differs from all aforementioned species by the absence of supranasals (present in Lygosoma) and having the first supralabial fused with nasal (not fused in Lygosma) (see Geissler et al. 2012). Sphenomorphus sheai also resembles Scincella apraefrontalis Nguyen, Nguyen, Böhme & Ziegler by having a low midbody scale count, no ear openings, and first supralabial fused with nasal; however, the new species can be distinguished from latter by the presence of prefrontals and opaque window on lower eyelid which are absent in Scincella apraefrontalis (Nguyen et al. 2010).