Goniurosaurus lichtenfelderi (Mocquard, 1897)

Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Le, Minh Duc, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, European Journal of Taxonomy 751 (1), pp. 38-67 : 52-54

Goniurosaurus lichtenfelderi (Mocquard, 1897)

Fig. 5

Diagnosis

Body robust; SVL 80.6−113.5 mm; external nares bordered by 5–10 nasal scales; supraorbital region with a row of slightly enlarged tubercles; outer surface of upper eyelid composed of granular scales, about one-half the size of those on top of head and without enlarged tubercles; internasals 1–5 (rarely 1:2; 2:2 or 2:3); supralabials 7–10; infralabials 6–9; preorbital scales 12–19; eyelid fringe scales 43–58; postmentals 2–6; paravertebral tubercles 22−33; scale rows around midbody 117–131, granular scales surrounding tubercles 10−13; axillary pockets shallow; subdigital lamellae under fourth toe 17−21; precloacal pores in males 25–33, in females 17−21; dorsal ground color of head, body and limbs dark purple-brown, without small dark brown blotches; transverse body bands 4, nuchal loop thin, posteriorly rounded, in U-shape; dorsal body bands between limb insertions 2, thin, light yellow; gular region and without dark spots; ventral surfaces of head, body and limbs dull white with a few dark dots on margin regions (modified after Grismer 2000; Grismer et al. 2002; Nguyen 2011).

Description (Supp. file 1: Table S2)

Body robust, adult males: SVL 80.6–113.5 mm (mean ± SE: 97.8 ± 0.7 mm, n =92), TaL 6.2– 84.8 mm (61.5± 1.7 mm); adult females: SVL 81.0– 105.5 mm (96.0± 0.7 mm, n=72), TaL 22.7– 81.2 mm (58.4 ± 1.4 mm); juveniles: SVL 41.1–77.1 mm (65.4± 2.9 mm, n=14), TaL 20.9–64.7 mm (48.3 ± 3.4 mm) (Supp. file 1: Table S2); head triangular, wider than neck, covered by uniform granular scales interspersed with tubercles in temporal and occipital regions; scales on rostrum slightly larger and flatter; enlarged supraorbital tubercles in a conspicuous row; middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial on each side, dorsolaterally by prenasal on each side, and dorsally by 1 or 2 internasals and two supranasals; internasals 1–5 (rarely 1:2; 2:2 or 2:3); external nares bordered by 5–10 nasals; preorbital scales 12–18; supralabials 7–10, grading into granular scales posteriorly; infralabials 6–9; eyes large, pupils vertical; eyelid fringe scales 47–58, those of upper eyelid slightly enlarged; outer surface of upper eyelid composed of granular scales of about one-half the size of those on top of head, without enlarged tubercles; fold of skin originating in the suborbital region extends posteroventrally across angle of jaw; external auditory meatus elliptical; tympanum deeply recessed; mental triangular, bordered laterally by first infralabial on each side and posteriorly by 2–5 postmentals; postmentals bordered by 7–10 gular scales; gular region below lower jaws without enlarged tubercles; gular scales juxtaposed and granular, abruptly grading posteriorly into flat hexagonal scales and even larger ventral scales.

Neck narrower than body, covered with uniform granular scales interspersed with several sharply conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat more flat; dorsal body tubercles surrounded by 10–13 granular scales; dorsal tubercle rows at midbody 15–22; paravertebral tubercles between limb insertions 22–33, distinct vertebral row of tubercles absent; scales around midbody 117–130; ventral scales large; males with 24–33 precloacal pores in a transverse continuous series, females without distinct precloacal pores (but 25–33 pitted scales present); region posterior to vent covered by flat juxtaposed scales and greatly swollen, containing 1 (rarely 2) enlarged tubercles on each side at the level of vent; tail thick at base; light band on tail base in width of 4–7 scales and with 6–10 tubercles in a transversal series.

Limbs robust, covered dorsally with granular scales interspersed with several tubercles and ventrally with flat juxtaposed to subimbricate scales; dorsal granular scales grade into slightly flattened subimbricate scales on top of pes and manus; hind limbs larger than forelimbs; axillary pockets shallow; subdigital lamellae wide, 6–9 under first finger, 13–17 under fourth finger, 6–10 under first toe, 16–20 under fourth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth (modified after Grismer 2000; Grismer et al. 2002; Nguyen 2011).

Coloration in life ( Fig. 5 )

Dorsal ground color of head, body and limbs grey and grey brown in juveniles without blotches and chestnut brown in adults with dark brown blotches (in animals with SVL> 80 mm); iris orange or redish brown; four bands on the dorsal ground, thin, yellow in both juveniles and adults, immaculate without dark spots (few dark brown spots present in some animals), all edged anteriorly and posteriorly by thin dark brown bands, including one thin nuchal loop extending from posterior corners of eyes and posteriorly rounded (in U-shape), two body bands between limb insertions, and another one on tail base; ground color of tail dark brown, and grey brown at mottled tail base; 3–5 immaculate white caudal bands except first band slightly yellow, and some animals with regenerated tail present immaculate grey brown without white caudal bands; ventral surfaces of head, body and limbs dull white with a few dark dots on margin regions of belly, limbs, gular region, but immaculate dull white in juveniles (SVL ≤ 65 mm) without dark spots.

Morphological comparisons

Our morphological comparisons of the tiger gecko species (except for G. araneus ) from Vietnam revealed an overall significant difference in the snout-vent length (SVL). As the result, G. huuliensis had the largest SVL, while G. lichtenfelderi had the shortest SVL in both adult males and females, compared to the other tiger geckos (P <0.05). However, the SVL of G. catbaensis was not significantly different from that of G. luii (P> 0.05, Supp. file 2: Fig. S1 ).

The PCA of 18 selected morphometric characters estimated the first (PC1) and second principal (PC2) components to explain 82.5% (74.5% and 8.0%, respectively) of the variance among the four investigated species of Goniurosaurus in Vietnam (Supp. file 2: Fig. S2 ). The extracted PC1 and PC2 scores of the PCA differ significantly among the four species (Kruskal-Wallis’s test, P <0.05), of which the values and morphometric spaces of G. lichtenfelderi are not concordant with the three remainders ( Fig. S2 ). Three morphological factors, namely head length (HL), head width (HW), and mouth width (MW) highly account for the overall difference in the PC1 score, while the PC2 score is mainly explained by the body height (BH) and body width (BW) (Supp. file 2: Fig. S2 ).

Regarding the comparisons of the 17 species of Goniurosaurus from China and Vietnam, the multiple correspondence analysis (MCA) clustered them separately into three groups ( G. lichtenfelderi group, G. luii group and G. yingdeensis group) represented with different meristic spaces on the spatial coordinate of Dim 1 and Dim 2 ( Fig. 6A , Table 1 ). The characters of fourth toe (LT4), body limb shape and body bands highly account for the Dim1 score, whereas the characters of precloacal pores (PP) and granular scales surrounding dorsal tubercles (GST) are the most correlated with the Dim 2 score ( Fig. 6B ).

Figures

Fig. 5. Goniurosaurus lichtenfelderi (Mocquard, 1897). A. Juvenile (SVL=55 mm). B. Juvenile (SVL=63 mm).C. Juvenile (SVL=72 mm).D. Adult (SVL=80 mm).E. Adult (SVL= 91 mm).F. Adult (SVL =97 mm).G. Adult (SVL= 105 mm).H. Adult (SVL=113 mm).

Fig. 1. Phylogram containing Goniurosaurus Grismer, Viets & Boyle, 1999 representatives from China, Japan and Vietnam based on the Bayesian analysis. Numbers above and below branches are MP/ML ultrafast bootstrap values and combined Bayesian posterior probabilities, respectively. Dash represents <50% value.

Fig. 2. Goniurosaurus catbaensis Ziegler, Nguyen, Schmitz, Stenke & Rösler, 2008. A. Juvenile (SVL=55 mm). B. Juvenile (SVL=69 mm). C. Juvenile (SVL=75 mm). D. Adult (SVL=91 mm). E.Adult (SVL=99 mm).F.Adult (SVL=105 mm).G. Adult (SVL=116 mm).H.Adult (SVL=125 mm).

Fig. 6. A. Multiple correspondence analysis (MCA) on meristic variation among 17 recorded species of Goniurosaurus Grismer, Viets & Boyle, 1999 in China and Vietnam. B. Correlation between meristic variables and principal dimensions (Dim1 and Dim2).