Cyrtodactylus peninsularis, Grismer & Kaatz & Grismer & Nguyen & Grergory & Wood Jr. & Murdoch & Anuar & Onn & Muin & Pawangkhanant & Suwannapoom & Poyarkov & Quah, 2025

Grismer, L. Lee, Kaatz, Amanda, Grismer, Jesse L., Nguyen, Eddie, Grergory, Jeren J., Wood Jr., Perry L., Murdoch, Matthew L., Anuar, Shahrul, Onn, Chan Kin, Muin, Muhamad A., Pawangkhanant, Parinya, Suwannapoom, Chatmongkon, Poyarkov, Nikolay A. & Quah, Evan S. H., 2025, The taxonomy of Cyrtodactylus consobrinus (Peters, 1871) (Squamata, Gekkonidae) and the description of a new species from the Thai-Malay Peninsula, ZooKeys 1241, pp. 105-137 : 105-137

Cyrtodactylus peninsularis sp. nov.

Figs 7 , 8 , 9

Gymnodactylus consobrinus — Boulenger 1912: 37 (part); Smith 1930: 13 (part); Bourret 2009: 123 (part).

Cyrtodactylus consobrinus — Grandison 1972: 81 (part); Dring 1979: 181 (part); Denzer and Manthey 1991: 314 (part); Kluge 2001: 8 (part); Cox et al. 1998: 88 (part); Chan-ard et al. 1999: 1058, 1064–65 (part), 2015: 49, 51 (part); Grismer 2008: 30 (part), 2011: 386 (part); Rösler 2016: 12 (part); Grismer and Quah 2019: 233; Sumarli et al. 2015: 5, 19 (part); Hong et al. 2021: 799, 800 & 807; Quah et al. 2021: 241 & 246; Davis et al. 2023: 3 (part); Poyarkov et al. 2023: 299, 301 (part).

Cyrtodactylus consubrinus — Manthey and Grossmann 1997: 221 (part; unjustified subsequent spelling).

Cyrtodactylus cf. consobrinus — Figueroa et al. 2023: 100 .

Cyrtodactylus (Cyrtodactylus) consobrinus — Rösler 2000: 65 (part).

Type material.

Holotype • Adult male ( LSUHC 10202 ) collected from the base of Gunung Belumut , Johor State, Peninsular Malaysia (2.065817°N, 103.526119°E at 245 m) on 8 September 2009 by Evan S. H. Quah, Perry L. Wood, Jr., Jesse L. Grismer, L. Lee Grismer, Shahrul Anuar, and Mohamed A. Muin . Paratypes. • Adult females: LSUHC 9333 northwestern lineage collected from Sungai Sedim , Kedah State, Peninsular Malaysia (5.414063°N, 100.779803°E at 129 m) on 16 March 2009 ; LSUHC 10048 northeastern lineage collected from Hutan Lipur Sekayu , Terengganu State, Peninsular Malaysia (4.980644°N, 102.934645°E at 441 m) on 27 March 2009 ; LSUHC 11136–37 north-central lineage collected from Gunung Stong , Kelantan State, Peninsular Malaysia (5.321880°N, 101.964944°E at 703 m) on 27 June 2009 ; LSUHC 11152 north-central lineage collected from Hutan Lipur Jelawang , Kelantan State, Peninsular Malaysia (5.340351°N, 101.969544°E at 699 m) on 27 June 2009 ; LSUHC 11979 northeastern lineage collected from Sungai Bubu , Terengganu State, Peninsular Malaysia (5.0120261°N, 102.952963°E at 77 m) on 1 September 2009 ; and LSUHC 12386 eastern lineage collected from Hutan Lipur Chemerong , Terengganu State, Peninsular Malaysia (4.660664°N, 103.001320°E at 129 m) on 17 August 2009 . Adult males: LSUHC 10230 southern lineage collected from the base of Gunung Belumut , Johor State, Peninsular Malaysia (2.045596°N, 103.530185°E at 244 m) on 9 September 2009 by Evan S. H. Quah, Perry L. Wood, Jr., Jesse L. Grismer, L. Lee Grismer, Shahrul Anuar, and Mohamed A. Muin ; and LSUHC 11267 northwestern lineage collected from trail 2, Sungai Enam , Belum, Perak State, Peninsular Malaysia (5.46768°N, 101.28961°E on 6 October 2009.

Additional specimens examine

(n = 52). See Suppl. material 5.

Diagnosis based on type series.

Cyrtodactylus peninsularis sp. nov. can be separated from all other species of the malayanus group by the combination of having a maximum SVL of 128.7 mm (female); 8–10 supralabials; 10–12 infralabials; 25–30 paravertebral tubercles; 15–20 rows of longitudinally arranged tubercles; 40–62 longitudinal rows of ventrals; 243–299 transverse rows of ventrals; 7–9 expanded subdigital lamellae on the fourth toe; 13–16 unmodified subdigital lamellae on the fourth toe; 21–25 total subdigital lamellae on the fourth toe; 21–25 total number of enlarged femorals; 2–9 total number of femoral pores in males, no femoral pores in females; 10–12 enlarged precloacals; nine or ten precloacal pores in males (n = 3), precloacal pores in some females (three of seven); two or three rows of large post-precloacals; two postcloacal tubercles (spines) on each side; dorsal pattern extremely variable, dark dorsal bands very wide reducing the pale dorsal interspaces to 2–4 thin lines; seven or eight dark and pale caudal bands (n = 3); large moderately keeled body tubercles; caudal tubercles extend beyond base of tail; subcaudals transversely expanded but not extending high up onto side of tail; enlarged distal femorals and enlarged precloacals not contiguous; no enlarged proximal femorals; top of head overlain with reticulating white network of thin lines; dark caudal bands wider than pale caudal bands; dark markings usually within pale caudal bands in adults (Tables 7 , 8 , Figs 5 , 6 , Suppl. material 2).

Description of holotype

(Fig. 7 , Table 7 ). Adult male SVL 113.1 mm; head moderate in length (HL / SVL 0.29), width (HW / HL 0.62), somewhat flattened (HD / HL 0.37), distinct from neck, triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; prefrontal deeply concave; canthus rostralis rounded; snout elongate (SN / HL 0.38), flat, rounded in dorsal profile; eye large (ED / HL 0.21); ear opening elliptical, obliquely oriented, moderate in size; eye to ear distance slightly greater than diameter of eye; rostral rectangular, partially divided dorsally, bordered posteriorly by large left and right supranasals and slightly smaller internasal, bordered laterally by first supralabials; external nares directed posterolaterally, bordered anteriorly by rostral, dorsally by large supranasal, posteriorly by six small postnasals, ventrally by first supralabial; nine (R, L) rectangular supralabials tapering to below posterior margin of eye, first six supralabials largest; 11 (R, L) infralabials tapering smoothly to slightly past termination of enlarged supralabials; scales of rostrum and lores raised, much larger than granular scales on top of head and occiput; scales of occiput intermixed with small, rounded, tubercles; superciliaries flat, elongate, largest dorsally; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapezoidal postmentals contacting medially for ~ 40 % of their length posterior to mental; one row of enlarged, sublabials extending posteriorly to fifth infralabials (R, L); gular and throat scales small, granular, grading posteriorly into slightly larger, flatter, smooth, subimbricate pectoral and ventral scales.

Body relatively long (AG / SVL 0.47) with well-defined ventrolateral folds; dorsal scales small, granular, interspersed with large, moderately keeled, semi-regularly arranged tubercles extending from occiput to beyond base of tail; ~ 16 longitudinal rows of tubercles at midbody; ~ 30 paravertebral tubercles; ~ 47 flat, imbricate, ventral scales much larger than dorsal scales; 12 enlarged precloacal scales not separated medially by poreless scales; no deep precloacal groove or depression; and three rows of large post-precloacal scales on midline.

Forelimbs moderate in length and stature (ForeL / SVL 0.16); granular scales of forelimbs slightly larger than those on body, large spinose tubercles on dorsal surface of forearms; palmar scales slightly rounded, juxtaposed; digits well-developed, inflected at basal interphalangeal joints, slightly narrower distal to inflections; subdigital lamellae transversely expanded, those proximal to joint inflections much wider than lamellae distal to inflections; claws well-developed, sheathed by a dorsal and ventral scale; hind limbs robust, wider and longer than forelimbs (TibL / SVL 0.18), covered dorsally by granular scales interspersed with large pointed tubercles; anterior scales of thigh slightly larger and flatter than dorsal scales of thigh; ventral scales of thighs rounded, subimbricate, slightly larger than dorsals; distal subtibials large, flat, subimbricate; one row of six (R, L) distal enlarged femoral scales, four on right bearing pores and three on left bearing pores, no other enlarged femoral scales; proximal femorals not forming an abrupt union with granular posteroventral scales of thigh; plantar scales rounded, juxtaposed; digits well-developed, inflected at basal interphalangeal joints; claws well-developed, sheathed by a dorsal and ventral scale at base; seven (R, L) wide subdigital lamellae on fourth toe proximal to joint inflection, 16 (R, L) narrower lamellae distal to joint inflection, 23 total subdigital lamellae.

Tail long (TL / SVL 1.21), original, tapering to a point; dorsal caudal scales small, generally square, juxtaposed; median row of subcaudals significantly larger than dorsal caudals, transversely expanded, not extending high up dorsally onto lateral side of tail; body tubercles extending beyond base of tail; hemipenial swellings at base of tail, two large postcloacal tubercles on both sides; and postcloacal scales flat, imbricate.

Color and pattern in preservative

(Fig. 7 ). No photograph of the living holotype was available. Ground color of top of head, limbs, and dorsum brown; top of head overlain with a reticulating network of thin white lines; snout bearing irregularly shaped white blotches and lines; thin, white, transverse line (i. e., interspace) on occiput and another immediately anterior to shoulders; a thin, white anterior dorsal line bifurcates paravertebrally forming two thin lines along anterior of flanks; a second thin white line occurs just posterior to midway between the limb insertions; another thin white line occurs at level of groin; all white lines bordered by large white tubercles and are thickly edged in dark brown; center of the brown regions between the thin white lines (i. e., body bands) bear irregularly shaped central pale brown areas; a thin white sacral line followed by seven widely spaced white caudal bands bearing darkened markings, separated by dark caudal bands nearly three times width of pale caudal bands; limbs dark brown to pale brown overlain with thin, white broken lines and irregularly shaped markings.

Etymology.

The species name peninsularis is in reference to the distribution of this species which is restricted to the Thai-Malay Peninsula of southern Thailand, Peninsular Malaysia, and Singapore.

Distribution.

Cyrtodactylus peninsularis sp. nov. ranges from extreme southern Thailand southward through nearly all habitats in Peninsular Malaysia to Singapore ( Grismer 2011) (Fig. 1 ). The Pulau Singkep population of Indonesia has not been investigated.

Variation.

Color pattern varies so extensively within Cyrtodactylus peninsularis sp. nov. it essentially defies a concise meaningful description (e. g., Fig. 8 A, C ). The type series was chosen to not only cover the geographic range of the species but to cover a great deal of its color pattern variation as well (Fig. 9 ). This remarkable variation, however, is paradoxically coupled to considerable conservatism in its morphological variation (Figs 4 , 5 , 6 , Tables 4 , 5 ).

Comparisons.

Cyrtodactylus peninsularis sp. nov. may be the sister species of C. consobrinus from which it differs morphometrically in having a statistically shorter snout-vent length; axilla-groin length; snout length; tibial and forelimb length; head length, width, and depth, and femur length (Table 3 ). It differs from C. consobrinus meristically by having statistically more infralabials; longitudinal rows of tubercles; enlarged precloacal scales; and fewer supralabial scales, longitudinal ventral scales, and paravertebral tubercles (Table 3 ). Hatchling C. peninsularis sp. nov. nearly always have thin white interspaces whereas they are yellow in C. consobrinus (Fig. 10 ). From the closely related C. hutan , C. peninsularis sp. nov. differs morphometrically in having a statistically shorter AG, SN, TibL, and ForeL, and shorter HL and HW. It differs meristically in having fewer SL, LVS, PS, and PVT and more IL and LRT (Table 3 ).

Natural history.

Much of the following is adapted from Grismer (2011). Cyrtodactylus peninsularis sp. nov. is a vagile, nocturnal, scansorial species that ranges throughout extreme southern Thailand, Peninsular Malaysia, and forested areas of Bukit Timah in Singapore up to ca 800 m in elevation ( Chan et al. 2019). Cyrtodactylus peninsularis sp. nov. is a relatively common inhabitant of primary and secondary dipterocarp forests and is occasionally found in peat swamps. Lizards are usually seen climbing on tree trunks, branches, exposed roots, and fallen logs where there are nearby crevices and holes into which they can quickly retreat when threatened. It is not uncommon to find multiple individuals on the same large tree up to 5 m above ground. Lizards are less commonly found among large boulders, taking refuge in cracks or in holes at their base. Anecdotally, this species seems to be more abundant in riparian areas. The holotype was found during the early evening at the base of Gunung Belumut at 245 m in elevation and 2 m above the ground on the trunk of a dipterocarp tree in primary forest (Fig. 11 ).

Figures

Figure 1. Maximum likelihood consensus topology based on 1459 base pairs of ND 2 and flanking tRNAs showing the relationships among the species of the malayanus group and among the lineages within Cyrtodactylus peninsularis sp. nov. Numbered individuals of C. peninsularis sp. nov. in the tree match the locality numbers on the distribution map of the six lineages on the Thai-Malay peninsula.

Figure 4. A MFA of the lineages of Cyrtodactylus peninsularis sp. nov. based on 15 morphometric and 16 meristic characters B percent distribution of the morphometric and meristic characters to the overall variation in Dims 1–4 C results of the PERMANOVA showing lineage pairs that differed significantly from each other.

Figure 5. A PCA of the lineages of Cyrtodactylus peninsularis sp. nov. based on 16 meristic characters B DAPC of the same data set based on the retention of the first three PCs C results of the PERMANOVA showing the lineages of C. peninsularis sp. nov. that plot significantly different from each other.

Figure 6. A PCA of the lineages of Cyrtodactylus peninsularis sp. nov. based on 15 morphometric characters B DAPC of the same data set based on the retention of the first two PCs C results of the PERMANOVA showing the lineages of C. peninsularis sp. nov. that plot significantly different from each other. The eastern lineage differs from all others.

Figure 7. A, C dorsal and ventral views of adult male holotype of Cyrtodactylus peninsularis sp. nov. LSUDPC 10202, respectively B gular region, throat, pectoral region, and ventral view of forelimbs D precloacal region, ventral view of hind limbs, and anterior subcaudal region.

Figure 8. Color and banding pattern variation among the lineages of Cyrtodactylus peninsularis sp. nov. A NEL — adult male, Lata Kekabu, Setiu, Terengganu, La Sierra University Digital Photograph Collection (LSUDPC) 13548, photo by Evan S. H. Quah B EL — juvenile, Endau-Rompin National Park, Johor, LSUHC 2585, photo by L. Lee Grismer C NWL — adult male, Sungai Enam, Perak LSUDPC 13549 (paratype LSUHC 11267), photo by Evan S. H. Quah D WL — adult female, Gunung Ledang, Johor LSUDPC 13550, photo by Evan S. H. Quah E NCL — adult female, Gunung Tebu, Terengganu LSUDPC 7997, photo by L. Lee Grismer F WL — adult female, Gunung Korbu, Perak, LSUDCP 13548, photo by Kin Onn Chan G NEL — adult male, Hutan Lipur Sekayu, Terengganu, LSUDPC 5951, photo by L. Lee Grismer H SL — adult female, Gunung Pulai, Johor, LSUDPC 13552, photo by Evan S. H. Quah.

Figure 9. Paratypes of Cyrtodactylus peninsularis sp. nov. showing the wide variation in dorsal banding pattern.

Figure 10. Cyrtodactylus consobrinus from Gua Angin, Sarawak, East Malaysia (Borneo) A adult female LSUDPC 10767 B adult male LSUDPC 10764 C hatchling LSUDPC 4928 D Juvenile LSUDPC 10769.

Figure 11. Habitat of the type locality at the base of Gunung Belumut, Johor State.