Morphological and molecular review of the Gekko diversity of Laos with descriptions of three new species Luu, Vinh Quang Calame, Thomas Nguyen, Truong Quang Le, Minh Duc Ziegler, Thomas Zootaxa 2015 3986 3 279 306 [151,458,462,488] Reptilia Gekkonidae Gekko Animalia Squamata 11 290 Chordata species sengchanthavongi sp. nov.     Holotype.VFU R.2014.14, adult male, in karst forest of Bualapha Town ( 17o19.444’N, 105o41.576’E, at an elevation of 210 ma.s.l.), Khammouane Province, central Laos, collected by Vinh Quang Luu and Thidtavanh Suliyavong on 20 June 2014.    Paratypes.Two adult males NUOLR-2015.3,  IEBRA.2015.33 and one adult female VFU R.2014.16, the same data as the holotype.   Diagnosis.  Gekko sengchanthavongi  sp. nov.can be distinguished from its congeners in the  Gekko japonicusgroup by a combination of the following characters: a medium-sized geckospecies (SVL 67.7–77.3 mm); nares in contact with rostral; internasals absent; postmentals enlarged; supralabials 8–10; infralabials 6 or 7; dorsal tubercles absent; ventral scale rows from mental to cloacal slit 175–184; scale rows around midbody 120–135; ventral scale rows 35–43; webbing weakly developed between fingers and toes; tubercles on dorsal surface of limbs and tail absent; precloacal pores 3+1 or 3+ 2 inthe males, separated from each other by one poreless scales, absent in the female; postcloacal tubercles 2/2; subcaudals enlarged; dorsum with dark bars.   Description of holotype.Adult male, total length 146.8 mm(SVL 67.7 mm, TaL 79.1 mm); body slender, elongate (ratio of AG/SVL 0.44); head longer than wide (ratio of HL/HW 1.48); rostral quadrangular, two times wider than high (ratio of RW/RH 2.14) and wider than mental (ratio of RW/MW 1.36 mm), without medial suture; rostral bordered by first supralabial and supranasal on each side; nostrils round, surrounded by rostral, first supralabial, supranasal, and two enlarged nasals posteriorly; posterior nasal region deeply concave; internasals absent; preorbital scales 17, preorbital region slightly concave; interorbital sacles 28; eye large (ratio of OD/HL 0.24), pupil vertical, superciliaries with six tiny spines posteriorly; ear oval–shaped, oblique, approximately half of eye diameter (ratio of TD/OD 0.48); mental triangular, wider than long (ratio of MW/ML 1.57); enlarged postmentals two, hexagonal, two times longer than wide, bordered by mental, first infralabial on each side and 6 gular scales posteriorly, outer gular scales larger than inner scales; supralabials 10/9; infralabials 7/7; dorsal scales smooth, small, round or oval, granular and juxtaposed; lateral skin folds present; ventral scales smooth, medial scales 3 or 4 times larger than dorsal scales, and largest in the middle of belly; ventral scales at midbody 35; scale rows around midbody 126; ventral scale rows between mental and cloacal slit 179; scales on dorsal limbs slightly enlarged; tubercles on limbs absent; enlarged femoral scales absent; fingers and toes webbed at base; subdigital lamellae under first finger 12/11, under fourth finger 14/13, under first toe 13/11, under fourth toe 15/15; precloacal pores 3+2, separated by one poreless scale, precloacal scales enlarged; postcloacal tubercles 2/2; tail basally thickened, without dorsal tubercles; subcaudals enlarged, smooth.  Coloration in life.Ground coloration of dorsal head dark brown; dorsal head and neck with dark and grey spots; dorsum with six bands between limb insertions, edged in yellowish brown, and irregular transverse dark brown bars; dorsal surface of limbs with dark blotches; dorsal surface of tail grey with 10 or 11 dark bands, edged in white; chin, throat, dorsolateral region and lower surface of limbs grey with light spots; chest, middle of belly grey; ventral tail grey in anterior part and with nearly closed bands in posterior part.  Sexual dimorphism.Measurements and scalation characters of the female paratypeare provided in Table 4. The following scale counts slightly vary between the female and males: scale rows in line from mental to the front of cloacal slit 175 ( 179–184 inmales), and precloacal pores absent in the female paratype.  Comparisons.Based on examination of specimens and data obtained from the literature ( Boulenger 1907; Ota et al.1995; Rösler et al.2005, 2010, 2011; Yang et al.2012, Nguyen et al.2013; Luu et al.2014; Ngo et al.2015; Yang 2015) we compared the new species from Laos with its congeners of the  Gekko japonicusgroup sensu Rösler et al.(2011)(see Table 5). Both correspondence and cluster analyses supported  Gekko sengchanthavongi  sp. nov.as a sister taxon to  G. bonkowskii  sp. nov.and  G. thakhekensis( Figs. 3−4). Molecular phylogenetic analyses also demonstrated the close relationships between these species (see Fig. 1).   FIGURE 7.A) Dorsal view and B) portrait of the holotype (VFU R.2014.14) of  Gekko sengchanthavongi  sp. nov.Photos V. Q. Luu.   Gekko sengchanthavongi  sp. nov.is lacking dorsal tubercles, which are present in the following species:  G. adleri,  G. auriverrucosus, G. c a n h i,  G. chinensis,  G. japonicus,  G. hokouensis,  G. kwangsiensis, G. liboensis,  G. palmatus,  G. scabridus,  G. shibatai,  G. similignum,  G. swinhonis,  G. vertebralis,  G. wenxianensis.   Gekko sengchanthavongi sp. nov.differs from  G. aaronbaueriby having fewer supralabials and infralabials (8–10 versus13 or 14 and 6 or 7 versus10 or 11, respectively), fewer interorbital scales (28–32 versus34–37), more scale rows around midbody (120–135 versus98–104), fewer lamellae under first toes (11–14 versus14–17), and more precoloacal pores in males (4 or 5 versus3 or 4); from  G. bonkowskii sp. nov.by a larger size (SVL reaching 77.3 versus69.2), having fewer supralabials and infralabials (8–10 versus12–14 and 6 or 7 versus10 or 11, respectively), more interorbitals (28–32 versus26 or 27), more scales from mental to cloacal slit (175–184 versus154–169), more scale rows around midbody (120–135 versus117), and fewer precloacal pores in males (4 or 5 versus6); from  G. melliby having a smaller size (SVL reaching 77.3 versus 84.6 mm), the absence of internasal ( versuspresent), postmentals enlarged ( versusnot enlarged), fewer infralabials (6 or 7 versus9–12), fewer scale rows around midbody (120–135 versus147–160), fewer ventral scales (35–43 versus43–49), and fewer precloacal pores in males (4 or 5 versus9–11); from  G. scientiadventuraby its larger size (SVL reaching 77.3 versus73.0 mm), having fewer interorbitals (28–32 versus41–51), more scales from mental to cloacal slit (175–184 versus118–140), fewer scale rows around midbody (120–135 versus139–143), and fewer precloacal pores in males (4 or 5 versus5–8); from  G. subpalmatusby the absence of internasals ( versuspresent), having fewer infralabials (6 or 7 versus7–12), postmentals enlarged ( versusnot enlarged), fewer ventral scales (35–43 versus48), and fewer precloacal pores in males (4 or 5 versus5–11); from G. t a i b a i e n s i sby its larger size (SVL reaching 77.3 versus69.0 mm), having fewer infralabials (6 or 7 versus8–10), having more lamellae under first and fourth toes (11–14 versus6 or 7 and 13–17 versus7 or 8, respectively); from  G. tawaensisby its larger size (SVL reaching 77.3 versus71.0 mm) the absence of internasals ( versuspresent), postmentals enlarged ( versusnot enlarged), and the presence of precloacal pores in males ( versusabsent); from  G. thakhekensisby having fewer infralabials (6 or 7 versus10 or 11), more ventral scales between mental and cloacal slit (175–184 versus165–174), more precloacal pores in males (6 versus1–5), more scale rows around midbody (120–135 versus110–116), and color pattern of dorsum (banded versusblotches); from  G. truongiby its smaller size (SVL reaching 77.3 versus 95.9 mm), having more ventral scales between mental and cloacal slit (175–184 versus160–172), and fewer precloacal pores in males (4 or 5 versus10–11); and from  G. yakuensisby the absence of internasals ( versuspresent), having postmentals enlarged ( versusnot enlarged), fewer precloacal pores in males (4 or 5 versus6–8), and the absence of dorsal tubercles on tail ( versuspresent).   Distribution.  G. sengchanthavongi  sp. nov.is currently known only from the typelocality in Bualapha Town, Khammouane Province, central Laos ( Fig. 5).   Etymology.We name the new species in honour of Mr. Sinnasone Sengchanthavong, Natural Resources and Environment Department of Khammouane Province, Laos, in recognition of his great support of our field research in Hin Nam No NPA. As common names, we suggest Sengchanthavong’s Gecko (English), Kap Ke Sengchanthavong (Laotian), and Sengchanthavongis Gecko (German).  TABLE 5.Morphological comparisons among the species of the  Gekko japonicusgroup (modified after Luu et al.2014, extended by Ngo et al.2015, Yang 2015, abbreviations defined in text, data unavailable). Character  G. bonkowskii G. aaronbaueri adleri auriverrucosus canhi chinensis japonicus hokouensis kwangsiensis  sp. nov.  sengchanthavongi  sp.nov. Maximum SVL (mm) 69.2 77.3 80.0 75.3 69 99.2 72 74 70 69.7 ......continued on the next pageContinued.    SPL (min) 12 8 13 10 9 14 10 9 10 10  SPL (max) 14 10 14 15 11 14 14 13 14 12  IFL (min) 10 6 10 9 9 10 9 8 8 11  IFL (max) 11 7 11 13 11 12 13 13 11 13  Nostril touching rostral 1 1 1 1 0 1 1 1 1 1  N (min) 3 3 3 3 3 3 2 3 3 3  N (max) 3 3 3 3 3 3 3 3 3 3  I (min) 0 0 0 1 0 1 1 0 1 0  I (max) 0 0 1 1 1 1 1 1 2 1  IO (min) 26 28 34 27 25 49 35 32 30 29  IO (max) 27 32 37 36 25 50 48 35 33 31  Postmentals 1 (enlarged = 1, not enlarged = 0) DTR (min) 0 1 0 1 0 1 7 0 16 1 1 11 10 0 9 0 12 1 9  DTR (max) 0 0 0 11 20 12 10 14 18 11  SMC (min) 154 175 - 168 - 168 156 169 153 185  SMC (max) 169 184 - 190 - 170 167 188 174 208  SR (min) 117 120 98 123 - 205 118 130 119 143  SR (max) 117 135 104 144 - 227 140 144 130 156  V (min) 37 35 39 35 - 49 37 39 36 41  V (max) 40 43 43 44 - 51 39 44 43 45    Character LT 1 (min)   G. bonkowskii  sp. nov.11   G. sengchanthavongi  sp.nov.11   aaronbaueri14   adleri11   auriverrucosus6   canhi13   chinensis8   japonicus10   hokouensis8   kwangsiensis11  LT1 (max) 13 14 17 14 8 16 10 12 11 13  LT 4 (min) 15 13 14 11 6 14 9 14 15 13  LT4 (max) 15 17 16 15 8 17 12 16 18 18  Toes webbed 1 1 1 0 0 1 0 0 1  Tubercles on fore-limbs (present = 1, absent = 0) Tubercles on hind limbs (present = 1, absent = 0) PP (in males, min) 0 0 6 0 0 3+1 0 0 3 0 1 17 1 1 8 0 1 5 0 1 17 1 1 6 0 0 5 0 0 9  PP (in males, max) 6 3+2 4 21 11 5 27 9 9 11  PAT (min) 0 2 1 1 2 2 1 2 1 1  PAT (max) 2 2 2 1 3 3 1 4 1 1  Tubercles on dorsal surface of tail (present = 1, absent = 0) Subcaudals enlarged 0 1 0 1 0 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1  Marking on upper head 0 0 1 1 0 0 0 0 0 0  Back flecked or blotched 1 1 1 0 0 1 1 0 1 0  Back banded 0 1 0 1 1 1 0 1 0 1  Tail banded 1 1 1 1 1 1 1 1 1 1  ......continued on the next page TABLE 5.(Continued) ......continued on the next pageContinued.    Character Maximum SVL (mm)  G. G. liboensis bonkowskii sengchanthavongi  sp. nov.sp.nov.69.2 77.3 85   melli84.6   palmatus79.7   scabridus77   scientiadventura shibatai73 70.9   similignum58.9   subpalmatus72  SPL (min) 12 8 12 10 11 9 12 10 12 8  SPL (max) 14 10 12 13 15 11 14 13 14 12  IFL (min) 10 6 11 9 9 9 9 10 11 7  IFL (max) 11 7 11 12 13 11 13 14 11 12  Nostril touching rostral 1 1 1 1 1 1 1 1 1 1  N (min) 3 3 3 3 3 3 3 3 3 3  N (max) 3 3 3 3 3 3 3 3 3 3  I (min) 0 0 0 1 0 1 0 0 1 1  I (max) 0 0 0 1 3 2 0 1 1 1  IO (min) 26 28 40 34 27 30 41 37 46 32  IO (max) 27 32 40 40 36 30 51 52 48 32  Postmentals (enlarged = 1, not enlarged = 0) DTR (min) 1 1 0 0 0 10 0 0 1 4 0 17 1 0 0 5 0 11 0 0  DTR (max) 0 0 10 0 12 21 0 14 11 0  SMC (min) 154 175 - 181 160 - 118 - - -  SMC (max) 169 184 - 200 194 - 140 - - -  SR (min) 117 120 - 147 116 - 139 114 144 -  SR (max) 117 135 - 160 147 - 143 134 153 -  V (min) 37 35 - 43 36 - 38 - - 48  V (max) 40 43 - 49 47 - 48 - - 48    Character LT 1 (min)   G. bonkowskii  sp. nov.11   G. sengchanthavongi  sp.nov.11   liboensis8   melli10   palmatus10   scabridus6   scientiadventura12    shibatai-   similignum11   subpalmatus7  LT1 (max) 13 14 8 12 13 9 15 - 13 9  LT 4 (min) 15 13 9 11 10 7 14 9 12 7  LT4 (max) 15 17 9 14 16 9 17 16 14 10  Toes webbed 1 1 0 1 1 0 1 0 1 1  Tubercles on fore-limbs (present = 1, absent = 0) Tubercles on hind limbs (present = 1, absent = 0) PP (in males, min) 0 0 6 0 0 3+1 0 0 - 0 0 9 0 0 23 1 1 10 0 0 5 0 0 0 0 0 17 0 0 5  PP (in males, max) 6 3+2 - 11 30 15 8 3 17 11  PAT (min) 0 2 1 1 1 1 2 1 1 1  PAT (max) 2 2 1 1 1 3 3 1 1 1  Tubercles on dorsal surface of tail (present = 1, absent = 0) Subcaudals enlarged 0 1 0 1 - - 0 1 1 1 1 1 0 1 1 1 1 1 0 1  Marking on upper head 0 0 0 1 0 0 1 0 0 0  Back flecked or blotched 1 1 0 0 1 1 1 1 1 1  Back banded 0 1 1 1 1 1 0 0 0 1  Tail banded 1 1 - 1 1 1 1 1 1 1  ......continued on the next page ......continued on the next pageContinued.     TABLE 5.(continued) Character Maximum SVL (mm)   G. bonkowskii  sp. nov.69.2   G. sengchanthavongi  sp.nov.77.3   swinhonis66   taibaiensis69   tawaensis71   thakhekensis79.2   truongi95.9   vertebralis69.2   wenxianensis59   yakuensis72  SPL(min) 12 8 7 9 13 12 13 10 12 12  SPL (max) 14 10 12 10 13 14 15 15 12 13  IFL(min) 10 6 7 8 15 10 11 10 11 9  IFL (max) 11 7 11 10 15 11 13 15 11 13  Nostril touching rostral 1 1 1 1 1 1 1 1 1 1  N (min) 3 3 - - - 3 3 3 - 3  N (max) 3 3 - - - 3 4 3 - 3  I (min) 0 0 - - 2 0 0 0 1 1  I (max) 0 0 - - 2 0 1 2 1 1  IO (min) 26 28 23 28 - 22 45 35 - -  IO (max) 27 32 24 28 - 26 48 50 - -  Postmentals (enlarged = 1, not enlarged = 0) DTR (min) 1 0 1 0 0 6 - - 0 0 1 0 1 0 0 2 - 10 0 -  DTR (max) 0 0 8 - 0 0 0 12 10 -  SMC(min) 154 175 - - - 165 160 - - -  SMC (max) 169 184 - - - 174 172 - - -  SR (min) 117 120 - - - 110 131 112 - -  SR (max) 117 135 - - - 116 143 139 - -  V (min) 37 35 40 - - 32 35 - 42 -  V (max) 40 43 40 - - 40 36 - 44 - Character G. G. swinhonis taibaiensis tawaensis thakhekensis truongi vertebralis wenxianensis yakuensis   bonkowskii sengchanthavongi  sp. nov. sp.nov.    LT 1 (min) 11 11 6 6 10 11 11 - 6 10  LT1(max) 13 14 9 7 10 13 13 - 6 10  LT 4 (min) 15 13 6 7 12 14 15 9 9 15  LT4(max) 15 17 9 8 12 15 17 17 9 15  Toeswebbed 1 1 0 - 0 1 0 0 0 0  Tubercles on fore-limbs 0 (present = 1, absent = 0) Tubercles on hind limbs 0 (present = 1, absent = 0) PP (in males, min) 6 0 0 3+1 1 1 7 - - 4 0 0 0 0 0 1 0 0 10 0 0 0 0 1 6 0 0 6  PP (in males, max) 6 3+2 9 6 0 5 11 1 8 8  PAT (min) 0 2 2 - 1 2 1 1 2 1  PAT (max) 2 2 3 - 1 2 1 2 3 1  Tubercles on dorsal surface of tail 0 (present = 1, absent = 0) Subcaudals enlarged 1 0 1 - 1 - - 0 1 0 1 0 1 0 1 - - 1 1  Marking on upper side of head 0 0 0 0 0 0 0 0 0 0  Back flecked or blotched 1 1 1 1 1 1 1 1 1 1  Backbanded 1 1 0 1 1 0 1 0 0 0  Tailbanded 1 1 1 1 1 1 1 - 1 1  Natural history.Specimens of  G. sengchanthavongiwere collected on karst walls at night from 20:00 to 21:30 during small rain, ca. 1.5–4 mabove the ground, at an elevation of ca. 210 ma.s.l. The surrounding area was disturbed secondary forest ( Fig. 8). 2465765987 2014-06-20 Khammouane Province VFU Vinh Quang Luu & Thidtavahn Suliyavong Laos 210 karst forest 17.324066 Bualapha Town 105.69293 11 290 VFU R.2014.14 1 1 holotype 2466413773 [199,736,673,698] 2014-06-20 Khammouane Province NUOL Vinh Quang Luu & Thidtavahn Suliyavong Laos 210 karst forest 17.324066 Bualapha Town 105.69293 11 290 NUOL R-2015.3 1 1 paratype 2466413768 [746,938,673,698] 2014-06-20 Khammouane Province IBER Vinh Quang Luu & Thidtavahn Suliyavong Laos 210 karst forest 17.324066 Bualapha Town 105.69293 11 290 IBER A.2015.33 1 1 paratype 2466413771 2014-06-20 Khammouane Province VFU Vinh Quang Luu & Thidtavahn Suliyavong Laos 210 karst forest 17.324066 Bualapha Town 105.69293 11 290 VFU R.2014.16 1 1 holotype