Morphological and molecular review of the Gekko diversity of Laos with descriptions of three new species Author Luu, Vinh Quang Author Calame, Thomas Author Nguyen, Truong Quang Author Le, Minh Duc Author Ziegler, Thomas text Zootaxa 2015 3986 3 279 306 journal article 10.11646/zootaxa.3986.3.2 13e6a51d-a4c4-4f79-8aae-9769050b4fce 1175-5326 234805 B07485D0-EE3A-4FE7-92E5-B6122240597B Gekko bonkowskii sp. nov . ( Fig. 2 ) Holotype . VFU R.2014.10, adult male, from the karst forest near Thakhek Town ( 17o27.260’N , 104o56.265’E , at an elevation of 146 m a.s.l.), Khammouane Province , central Laos , collected by Vinh Quang Luu and Thomas Calame on 6 June 2014 . Paratype . NUOL R-2015.1, adult female, the same data as the holotype . Diagnosis. Gekko bonkowskii sp. nov. differs from its relatives by a combination of the following characters: a medium-sized gecko species (SVL 66.7 mm in the male, 69.2 mm in the female); nares touching rostral; internasals absent; postmentals enlarged; dorsal tubercles absent; ventral scale rows from mental to cloacal slit 154–169; scale rows around midbody 110–116; ventral scale rows 37–40; webbing weakly developed between fingers and toes; dorsal surface of limbs and tail without tubercles; precloacal pores six in a continuous row in the male, absent in the female; postcloacal tubercles 0–2; subcaudals enlarged; dorsum with black and grey blotches. FIGURE 2. A) Dorsal view of the holotype (VFU R.2014.10) and B) the paratype (NUOL R–2014.5) of Gekko bonkowskii sp. nov. Photos T. Calame. Description of holotype . An adult male with a total length of 145.9 mm (SVL 66.6 mm , TaL 79.3 mm ); body slender, elongate (ratio of AG/SVL 0.48); head longer than wide (ratio of HL/HW 1.43); rostral quadrangular without suture medially, nearly twice wider than high (ratio of RW/RH 1.93) and wider than mental (ratio of RW/ MW 1.29 mm ), touching first supralabial and supranasal on each side; nostrils round, in contact with rostral, first supralabial, supranasal, and two enlarged nasals posteriorly, upper nasal smaller than lower nasal; posterior nasal region concave; internasal absent; preorbitals 24, preorbital region deeply concave; interorbitals 26; eye large (ratio of OD/HL 0.23), pupil vertical; ear opening oval, oblique, smaller than eye (TD/OD ratio 0.38); mental triangular, about two times wider than long (ratio of MW/ML 1.93); postmentals two, hexagonal, twice as long as wide, and longer than length of mental, touching mental, first infralabial on both sides and 6 gular scales posteriorly, outer gular scales larger than inner scales; supralabials 12/12; infralabials 10/11; dorsal scales on body smooth, round or oval, granular and juxtaposed; lateral fold present; ventrals distinctly larger than dorsal scales, smooth, imbricate, and largest in the middle of belly; ventral scale rows at midbody 37; scale rows around midbody 117; ventral scales in a row between mental and cloacal slit 154; scales on dorsal 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 finger 12/12, under fourth finger 14/13, under first toe 12/12, under fourth toe 15/15; precloacal pores six, in a continuous row; precloacal scales enlarged; postcloacal tubercles 2/0; base of tail thickened, without tubercles on dorsal surface; subcaudals enlarged, smooth, imbricate. Coloration in life. Dorsal surface of head brownish grey with dark and grey blotches; labials with grey bars; upper eyelids brownish black; dorsal surface of body brownish grey with black and grey blotches, in oval or round shape, forming a cross-row in anterior part, irregular in posterior part; dorsal surface of fore and hind limbs dark grey; ventral surface of head, belly, and limbs cream with black dots, chest cream with black blotches on each scale; dorsal surface of tail with six or seven grey transverse bands; ventral tail grey in forepart and with nearly closed bands in hindpart. Sexual dimorphism. Measurements and scalation characters of the female paratype are shown in Table 4 . The following scale counts slightly vary between the paratype and the holotype : scale rows from mental to the front of cloacal slit 154 ( 169 in the holotype ), ventrals 37 ( 40 in the holotype ), and precloacal pores absent in the female paratype . Throat and flanks of the paratype reticulated. 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 the remaining members of the Gekko japonicus group sensu Rösler et al. (2011) (see Table 5). Both correspondence and cluster analyses revealed that Gekko bonkowskii sp. nov. is closely related to G. thakhekensis ( Figs. 3−4 ). Molecular phylogenetic analyses supported the sister relationship between the new species and G. thakhekensis (see Fig. 1 ). Morphologically, the new Gekko species can be distinguished from the species of the G. japonicus (following Rösler et al. 2011 ) group as follows: FIGURE 3. Correspondence analysis showing species association of the Gekko japonicus group sensu Rösler et al. (2011) based on morphological comparisons. TABLE 4. Measurements (in mm) and morphological characters of the type series of Gekko bonkowskii sp. nov. and Gekko sengchanthavongi sp. nov. (for abbreviations see material and methods). Gekko bonkowskii sp. nov. Gekko sengchanthavongi sp. nov.

VFU R.2014.10 (holotype)	NUOL R- 2014.5 (paratype)	VFU R.2014.14 (holotype)	IEBR A.2015.33 (paratype)	NUOL R- 2015.3 (paratype)	Min–Max	M±SD	VFU R.2014.13 (paratype)
Sex	adult male	adult female	adult male	adult male	adult male	adult female
SVL TaL	66.7 79.2	69.2 76.0	67.7 79.1	77.3 8.0*	72.8 80.0*	67.7–77.3 79.1(n=1)	72.6±4.8 79.1±0.0	76.8 84.6
AG	31.8	31.5	29.6	32.2	29.4	29.4–32.2	30.4±1.6	35.5
HL	18.5	18.4	19.8	21.6	21	19.8–21.6	20.8±0.9	21.6
HW	12.9	12.8	13.4	14.3	17.3	13.4–17.3	15.0±2.0	13.8
HH	7.6	7.3	7.4	7.9	8.5	7.4–8.5	7.9±0.6	7.7
SE	8.5	7.7	8.6	8.4	8.9	8.4–8.9	8.6±0.3	9.2
OD	4.2	4.3	4.8	4.8	4.7	4.7–4.8	4.8±0.1	5.2
TD	1.6	1.7	2.3	2.2	2.3	2.2–2.3	2.3±0.1	2.2
RW	2.7	2.9	3.0	3.2	3.3	3.0–3.3	3.2±0.2	3.3
RH	1.4	1.4	1.4	1.2	1.5	1.2–1.5	1.4±0.2	1.4
MW	2.1	1.8	2.2	2.3	2.3	2.2–2.3	2.3±0.1	2.4
ML	1.5	1.6	1.4	1.8	1.4	1.4–1.8	1.5±0.2	1.7
CS	4/5	4/3	6/6	5/6	5/5	5–6	5.5±0.6	5/6
N	3/3	3/3	3/3	3/3	3/3	3	3.0±0.0	3/3
I	0	0	0	0	0	0	0	0
SPL	12/12	14/12	10/9	10/8	8/9	8–10	9.0±0.9	9/10
IFL	10/11	10/10	7/7	7/7	6/7	6–7	6.8±0.4	7/7
PO	24	24	17/17	18/18	18/18	17–18	17.7±0.5	18/18
IO	26	27	28	30	32	28–32	30.0±2.0	32
PM	2	2	2	2	2	2	2.0±0.0	2
GP	6	6	6	7	6	6–7	6.3±0.6	7
DTR	0	0	0	0	0	0	0	0
GSDT	0	0	0	0	0	0	0	0
SMC	154	169	179	184	182	179–184	181.7±2.5	175
SR	117	117	126	120	135	120–135	127.0±7.6	130
V	37	40	35	40	43	35–43	39.3±4.0	42
LF1	12/12	10/10	12/11	11/10	15/13	10–15	12.0±1.8	12/11
LF4	14/13	13/14	14/13	13/12	15/15	12–15	13.7±1.2	13/13
LT1	12/12	13/11	13/11	12/11	13/14	11–14	12.3±1.2	13/13
LT4	15/15	15/15	14/15	14/13	17/16	13–17	14.8±1.5	15/15
PP	6	0	3+2	3+2	3+1	4–5	4.7±0.6	0
PAT	2/0	2/0	2/2	2/2	2/2	2	2.0±0.0	2/2

G. bonkowskii sp. nov. can be distinguished from G. aaronbaueri Ngo, Pham, Phimvohan, David & Teynié by its smaller size (SVL reaching 69.2 versus 80.0 mm), having fewer interobital scales (26–27 versus 34–37), more scale rows around midbody (117 versus 98–104), fewer subdigital lamellae under first toe (11–13 versus 14–17), and more precloacal pores in males (6 versus 3–4); from G. adleri by the absence of internasals, tubercles on dorsal surface of limbs and tail ( versus present), having fewer scale rows around midbody (117 versus 123–144) and fewer precloacal pores in males (6 versus 17–21); from G. auriverrucosus Zhou & Liu by having more supralabials (12–14 versus 9–11), nostril touching rostral ( versus not touching), postmentals enlarged ( versus not enlarged), lacking dorsal tubercles ( versus present), fewer precloacal pores in males (6 versus 8–11), and fewer postcloacal tubercles (0–2 versus 2–3); from G. c a n h i by having a smaller size (SVL reaching 69.2 mm versus 99.2 mm ), the absence of internasals and dorsal tubercles ( versus present), fewer scale rows around midbody (117 versus 205– 227), fewer ventral scales (37–40 versus 49–51), and the absence of tubercles on dorsal surface of hind limbs ( versus present); from G. ch i n e n s i s Gray by the absence of internasals ( versus present), fewer interorbital scale rows (26–27 versus 35–48), the absence of tubercles on dorsal surface of body, hind limbs, and tail ( versus present), and fewer precloacal pores in males (6 versus 17–27); from G. japonicus (Schlegel) by having postmentals enlarged ( versus not enlarged), the absence of tubercles on dorsal surface of body, limbs, and tail ( versus present), fewer scales around midbody (117 versus 130–144), fewer precloacal pores in males (6 versus 6– 9), and fewer postcloacal tubercles (0–2 versus 2–4); from G. hokouensis Pope by the absence of internasals and dorsal tubercles ( versus present), fewer interorbitals (26–27 versus 30–33), postmentals enlarged ( versus not enlarged), and the absence of tubercles on dorsal surface of tail ( versus present); from G. kwangsiensis Yang by the absence of tubercles on dorsal surface of body ( versus present in G. kwangsiensis ), fewer scales from mental to cloacal slit (154–169 versus 185–208), fewer ventral scales (37–40 versus 41–45), and fewer precloacal pores in males (6 versus 9–11); from G. liboensis Zhao & Li by its smaller size (SVL reaching 69.2 versus 85.0 mm), having fewer interorbitals (26–27 versus 40), and the absence of tubercles on dorsal surface of body ( versus present); from G. melli Vogt by its smaller size (SVL reaching 69.2 versus 84.6 mm ), internasals absent ( versus present), postmentals enlarged ( versus not enlarged), fewer scales from mental to cloacal slit (154–169 versus 181–200), fewer scale rows around midbody (117 versus 147–160), fewer ventral scale rows (37–40 versus 43–49), and fewer precloacal pores in males (6 versus 9–11); from G. palmatus by its smaller size (SVL reaching 69.2 versus 79.7 mm ), the absence of tubercles on dorsal surface of body and tail ( versus present), and fewer precloacal pores in males (6 versus 23–30); from G. scabridus Liu & Zhou by the absence of internasals ( versus present), postmentals enlarged ( versus not enlarged), the absence of tubercle on dorsal surface of body, limbs, and tail ( versus present), and fewer precloacal pores in males (6 versus 10–15); from G. scientiadventura by having fewer interorbitals (26– 27 versus 41–51), more scales from mental to cloacal slit (154–169 versus 118–140), and fewer scale rows around midbody (117 versus 139–143); from G. shibatai Toda, Sengoku, Hikida & Ota by having fewer interorbitals (26– 27 versus 37–52), postmentals enlarged ( versus not enlarged), the presence of toe webbing ( versus absent), the absence of tubercles on dorsal surface of body and tail ( versus present), and more precloacal pores (6 versus 0–3); from G. similignum Smith by its larger size (SVL reaching 69.2 versus 58.9 mm ), having fewer interorbitals (26–27 versus 46–48), the absence of internasals ( versus present), postmentals enlarged ( versus not enlarged), the absence of tubercles on dorsal surface of body and tail ( versus present), fewer scale rows around midbody (117 versus 144– 153), and fewer precloacal pores in males (6 versus 17); from G. subpalmatus Günther by having fewer interorbitals (26–27 versus 32), postmentals enlarged ( versus not enlarged), the absence of internasals ( versus present), and fewer ventral scales (37–40 versus 48); from G. swinhonis Günther by having postmentals enlarged ( versus not enlarged), the absence of dorsal tubercle rows on body and limbs ( versus present), fewer precloacal pores in males (6 versus 7–9), and fewer postcloacal tubercles (0–2 versus 2–3); from G. t a i b a i e n s i s Song by having more lamellae under first and fourth toes (11–13 versus 6−7 and 15 versus 7 or 8, respectively) and more precloacal pores in males ( 6 in a continous series versus 4–6 in interrupted series); from G. tawaensis Okada by the lack of internasals ( versus 2), postmentals enlarged ( versus not enlarged), and precloacal pores present ( versus absent); from G. thakhekensis by its smaller size (SVL reaching 69.2 versus 79.2 mm ), having toe webbing developed only basally (ca. one seventh) ( versus one fifth of length of digits), and more precloacal pores in males (6 versus 1–5); from G. truongi by its smaller size (SVL reaching 69.2 versus 95.9 mm ), having fewer scale rows around midbody (117 versus 131–143), and fewer precloacal pores in males (6 versus 10–11); from G. vertebralis Toda, Sengoku, Hikida & Ota by having fewer interorbitals (26–27 versus 35−50), postmentals enlarged ( versus not enlarged), and the absence of tubercles on dorsal surface of body and tail ( versus present); from G. wenxianensis Zhou & Wang by its larger size (SVL reaching 69.2 versus 59.0 mm), the absence of internasals ( versus present), the absence of tubercles on dorsal surface of body and hind limbs ( versus present), fewer ventral scale rows (37–40 versus 42−44), and fewer precloacal pores in males (6 versus 6–8); and from G. yakuensis Matsui & Okada by lacking internasals ( versus present), having postmentals enlarged ( versus not enlarged), fewer precloacal pores in males (6 versus 6–8), and the absence of tubercles on dorsal surface of tail ( versus present). FIGURE 4. Cluster analysis showing species correlation of the Gekko japonicus group based on morphological comparisons (1000 bootstrap replicates). Distribution. Gekko bonkowskii sp. nov. is currently known only from the type locality in Khammouane Province, central Laos ( Fig. 5 ). Etymology. The new Gekko species is named after Professor Dr. Michael Bonkowski from the Zoological Institute, University of Cologne, Germany to acknowledge his engagement for herpetological and ecological research in the Indochina region. We suggest as common names: Bonkowski’s Gecko (English) , Kap Ke Bonkowski (Laotian), and Bonkowskis Gecko (German) . FIGURE 5. Map showing the type locality (black circle) of three new species of Gekko in Khammouane Province, central Laos. FIGURE 6. A) Agricultural area and macrohabitat of Gekko bonkowskii sp. nov. , B) construction works, and C) limestone exploitation for building purposes Photos V. Q. Luu. Natural history. Specimens of Gekko bonkowskii were found at night between 20:00 and 21:00 on the tree trunk of shrubs, about 1.0– 1.5 m above the ground, near the entrance of a karst cave at an elevation of 146 m a.s.l. Surrounding habitat was secondary forest of small hardwood and shrubs near a village (ca. 20 m ) and about 40 m from the main road. The crepuscular or nocturnal new species co-occurs with at least two other gecko species in the same karstic microhabitat: Gekko gecko and the recently described bent-toed gecko Cyrtodactylus jaegeri (Luu et al. 2014) . We also found the large huntsman spider species Heteropoda maxima (Jaeger) in the immediate vicinity of the observed gecko species ( Fig. 6 ).