Cyrtodactylus macrotuberculatus Grismer and Ahmad, 2008

Termprayoon, Korkhwan, Rujirawan, Attapol, Grismer, L. Lee, Jr., Perry L. Wood & Aowphol, Anchalee, 2021, Taxonomic reassessment and phylogenetic placement of Cyrtodactylus phuketensis (Reptilia, Gekkonidae) based on morphological and molecular evidence, ZooKeys 1040, pp. 91-121 : 91

Cyrtodactylus macrotuberculatus Grismer and Ahmad, 2008 Figure 4

Cyrtodactylus macrotuberculatus Grismer & Ahmad, 2008: 55; Grismer 2011: 406; Grismer et al. 2012: 45.

Cyrtodactylus phuketensis Sumontha et al., 2012: 62.

Type specimens.

Holotype (adult male, ZRC 2.6754) from Malaysia, Kedah, Pulau Langkawi, Gunung Raya; Paratypes: Malaysia, Kedah, Pulau Langkawi, Gunung Raya: ZRC 2.6755-2.6756, Telaga Tujuh: ZRC 2.6757, Lubuk Semilang: ZRC 2.6758.

Additional specimens examined

(including types of C. phuketensis ). Malaysia - Kedah, Pulau Langkawi , Gunung Raya: LSUHC 09428-09429, LSUHC 09432 ; Perlis, Perlis State Park : LSUHC 09981, LSUHC 10097, ZRC 2.4869. Thailand - Satun Province, Mueang Satun District, Adang Island : ZMKU R 00871 - 00875, ZMKU R 00879 - 00882, Rawi Island : ZMKU R 00883 -00889; Songkhla Province, Hat Yai District , Chalung Sub-district: ZMKU R 00876 -00878; Phuket Province, Thalang District : PSUZC-RT 2010.58, THNHM 15378, ZMKU R 00894 -00896, Kathu District: ZMKU R 00890 - 00893, ZMKU R 00897 - 00898 ( Table 1 ) .

Expanded diagnosis.

Cyrtodactylus macrotuberculatus can be separated from all other species of C. pulchellus complex by having the following combination of characters (Table 6 ): (1) maximum SVL 117.9 mm (mean 105.0 ± SD 9.8, N = 39); (2) 9-13 supralabial and 7-11 infralabial scales; (3) prominent tuberculation on body; (4) tubercles on ventral surface of forelimbs, gular region, in ventrolateral body folds, and anterior one-third of tail; (5) 34-49 paravertebral tubercles; (6) 19-27 longitudinal tubercle rows; (7) 17-28 ventral scales; (8) 19-23 subdigital lamellae on the fourth toe; (9) 28-42 femoroprecloacal pores in males; (10) deep precloacal groove in males; (11) three or four dark dorsal body bands; (12) body band wider than interspace; (13) 7-10 (N = 12) ringed dark caudal bands on original tail; (14) white caudal bands infused with dark pigmentation in adults; (15) posterior portion of tail in hatchlings and juveniles bands not white.

Description of adult males.

SVL of adult males range from 88.9-117.9 mm (mean 105.7, N = 24); head moderate in length (HL/SVL 0.27-0.30), width (HW/HL 0.62-0.74), somewhat flattened (HD/HL 0.37-0.45), distinct from neck, triangular in dorsal profile; lores concave; frontal and prefrontal regions deeply concave; canthus rostralis sharply rounded; snout elongate (ES/HL 0.37-0.41), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.21-0.27); ear opening elliptical, moderate in size (EL/HL 0.05-0.10) obliquely oriented; eye to ear distance greater than diameter of eye; rostral rectangular, divided dorsally by an inverted Y or I-shaped furrow, bordered posteriorly by large left and right supranasals and small internasal, bordered laterally by external nares and first supralabials; external nares bordered anteriorly by rostral, dorsally by one large anterior supranasal, posteriorly by two postnasals, ventrally by first supralabial; 9-13 rectangular supralabials extending to just beyond upturn of labial margin, tapering abruptly below midpoint of eye; 7-11 infralabials not tapering in size posteriorly; scales of rostrum and lores slightly raised, larger than granular scales on top of head and occiput, those on posterior portion of canthus rostralis slightly larger; scales on top of head and occiput intermixed with enlarged tubercles; large, boney frontal ridges bordering orbit confluent with boney, transverse, parietal ridge; dorsal superciliaries elongate, smooth, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by left and right trapezoidal postmentals that contact medially for 40-50% of their length posterior to mental; single row of slightly enlarged, elongate sublabials extending posteriorly to 5th-7th infralabial; small, granular, gular scales intermixed with numerous large, conical tubercles grading posteriorly into larger, conical tubercles on throat which abruptly transition into large, flat, smooth, imbricate, pectoral and ventral scales.

Body relatively short (AG/SVL 0.43-0.51) with well-defined, tuberculate, ventrolateral folds; dorsal scales small, granular, interspersed with large, trihedral, regularly arranged, keeled tubercles separated by no more than three granules at their base; tubercles extend from top of head onto approximately one-half of tail but not onto regenerated tail; tubercles on occiput and nape relatively small, those on body largest; approximately 19-27 longitudinal rows of dorsal tubercles at the mid body; approximately 37-49 paravertebral tubercles; 17-28 flat, imbricate, ventral scales and much larger than dorsal scales; precloacal scales large, smooth; deep precloacal groove (= depression).

Forelimbs moderate in stature, relatively short (FL/SVL 0.15-0.17); virtually no granular scales on dorsal surface of forelimbs, only large, trihedral, keeled tubercles; palmar scales slightly rounded; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae nearly square proximal to joint inflection, only slightly expanded distal to inflection; digits more narrow distal to joints; claws well-developed, sheathed by dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.18-0.21), virtually no granular scales on dorsal surfaces of hind limbs, only large, trihedral, keeled tubercles; ventral scales of thigh flat, smooth, imbricate; ventral, tibial scales flat, imbricate, slightly keeled; two rows of enlarged, flat, imbricate, femoroprecloacal scales extend from knee to knee through precloacal region where they are continuous with enlarged, pore-bearing precloacal scales; 28-42 contiguous, pore-bearing femoroprecloacal scales forming an inverted T bearing a deep, precloacal groove; eight to eleven pores bordering groove; postfemoral scales immediately posterior to the pore-bearing scale row conical, forming an abrupt union on posteroventral margin of thigh; plantar scales low, slightly rounded; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae proximal to joint inflection nearly square, only slightly expanded distal to inflection; digits more narrow distal to joints; claws well-developed, sheathed by a dorsal and ventral scale; 19-23 subdigital lamellae on the 4th toe.

Original tail (TL/SVL) moderate in proportions, 123-135% of SVL (mean 128, N = 12), 7.9-11.6 mm in width at base, tapering to a point; dorsal scales at base of tail square, smooth, flat, subimbricate, lacking tubercle on regenerated tail; median row of transversely enlarged, subcaudal scales; shallow caudal furrow; two to five small, postcloacal tubercles at base of tail on hemipenial swellings; all postcloacal scales flat, large, imbricate.

Coloration of adult male ZMKU R 00871 in life

(Fig. 5 ). Ground color of head, body, limbs, and dorsum light-brown to yellowish brown; wide, dark-brown nuchal band edged anteriorly and posteriorly by thin, creamy-white lines bearing tubercles extends from posterior margin of one eye to posterior margin of other eye; four similar body bands between nuchal loop and hind limb insertions edged anteriorly and posteriorly by thin, creamy-white lines bearing tubercles, first band terminates at shoulders, second and third bands terminate just dorsal to ventrolateral folds, the fourth band terminates at femurs; dark body bands slightly larger than light-colored interspaces; one additional dark-brown band posterior to hind limbs; original portion of tail bearing eight ringed, dark-colored bands separated by seven, narrower, off-white bands infused with dark pigmentation; ventral surfaces of head smudged with brown; abdomen and limbs beige, slightly darker, lateral regions.

Coloration in preservative

(Fig. 6 ). Color pattern of head, body, limbs, and tail similar to that in life with some fading. Ground color of head, body, limbs, and dorsum tan; dark body and dark caudal bands lighter than in life.

Variation.

Cyrtodactylus macrotuberculatus usually varies in coloration and banding pattern (Figs 7 - 8 ). In females, a precloacal groove and pores are absent (Fig. 9 ). PSUZC-RT 2010.58 and THNHM 15378 have a shallow precloacal groove. Three dark body bands occur in PSU 2010.58, THNHM 15378, ZMKU R 00889-00894 and ZMKU R 00897. In ZMKU R 00887, the second dorsal band bifurcates just dorsal to the ventrolateral fold. ZMKU R 00895 has four bands and the third band is incomplete. The third body band in ZMKU R 00896 is broken on the left of the midline and contacts the fourth body band bilaterally. Nuchal loop and body bands of ZMKU R 00883, ZMKU R 00895, and ZMKU R 00898 edged anteriorly and posteriorly by thin, light-yellow lines and tubercles; and dorsal superciliaries are light-yellow (Fig. 8 ). Variation in morphometric and meristic data are shown in Table 6 .

Distribution.

Cyrtodactylus macrotuberculatus is distributed on the mainland and only known from one island in Peninsular Malaysia and southern Thailand (Fig. 1 ). This species is known from Pulau Langkawi (Gunung Raya, Telaga Tujuh, Gunung Machinchang, and Lubuk Semilang), Kedah, Peninsular Malaysia ( Grismer and Ahmad 2008). Other populations are found from Peninsular Malaysia; Kedah (Bukit Wang, Gunung Jerai, Hutan Lipur Sungai Tupah, Kuala Nerang, and Ulu Muda) and Perlis (Bukit Chabang, Chuping and Perlis State Park; [ Grismer 2011; Grismer et al. 2012; Quah et al. 2019]). In Thailand, C. macrotuberculatus was recorded from Phatthalung Province ( Grismer et al. 2012); Phuket Province, Kathu District (Kathu Waterfall) and Thalang District (Thep Krasatti Sub-district, previously type locality of C. phuketensis ); Satun Province, La-ngu District, and Mueang Satun District (Adang and Rawi Islands); Songkhla Province, Rattaphum District ( Grismer et al. 2012) and Hat Yai District (Ton Nga Chang Waterfall).

Natural history.

Based on specimens in Thailand, all individuals were found in similar habitat type, lowland forest habitat along granitic rock streams and surrounding areas (elevation 7-186 m asl) during a night survey (1900-2200; Fig. 10 ). The geckos were found mostly on rock boulders, vegetation (trunk of tree, buttress root, rotting wood and vines), and sometimes on the ground with leaf litter and high humidity (26.3-30.8 °C in temperature, 73.8-100% in relative humidity). Gravid female (ZMKU R 00876) contained four eggs during December. One juvenile (ZMKU R 00898, 56.50 mm in SVL) was found on a tree trunk in January. The varied microhabitats within which this species occurs, are consistent with its characterization as a habitat generalist ( Grismer et al. 2020, 2021b) and may account for its wide peninsular and insular distribution relative to other species of the Cyrtodactylus pulchellus group whose distributions are much less extensive or site-specific ( Grismer et al. 2012, 2014, 2016; Quah et al. 2019; Wood et al. 2020).

In Thailand, C. macrotuberculatus were found sympatric with other gecko species, Cnemaspis adangrawi Ampai et al., 2019 on Adang and Rawi Islands, Satun Province ( Ampai et al. 2019); Cnemaspis phuketensis Das and Leong, 2004, Cyrtodactylus oldhami Theobald, 1876, and Gekko (Ptychozoon) tokehos Grismer et al., 2019 at Kathu and Thalang District, Phuket Province; G. (P.) tokehos , Cnemaspis kumpoli Taylor, 1963, and Gehyra mutilata (Weigmann, 1834) at Hat Yai District, Songkhla Province.

Comparison.

Cyrtodactylus macrotuberculatus is distinguished from all other 15 species in the C. pulchellus complex by a combination of morphological characters (Table 7 ). It differs from all other species by having prominent tuberculation on the body; tubercles on ventral surface of forelimbs, gular region, and in ventrolateral body folds; 34-49 paravertebral tubercles; 19-27 longitudinal tubercle rows; 17-28 ventral scales; 19-23 subdigital lamellae on the fourth toe; 28-42 femorprecloacal pores in males; deep precloacal groove in males; no scattered white spots on dorsum; 7-10 dark-ringed caudal bands on original tail; white caudal bands on original tail infused with dark pigmentation in adults. Additional comparisons between C. macrotuberculatus and other species in C. pulchellus complex are in Table 7 .

Based on molecular data, C. macrotuberculatus is the sister lineage to a clade composed of C. pulchellus and C. evanquahi . It can be separated from those two species by having tubercles on ventral surface of forelimbs, gular region, and in ventrolateral body folds (vs. absent in C. evanquahi and C. pulchellus ); 17-28 ventral scales (vs. 29-33 in C. evanquahi and 29-34 in C. pulchellus ); deep precloacal groove in males (vs. a shallow in C. evanquahi ); three or four dark dorsal bands (vs. six or seven bands in C. evanquahi and only four bands in C. pulchellus ); white posterior caudal region absent (vs. present in C. evanquahi ); hatchlings and juveniles without white tail tip (vs. present in C. evanquahi ).

Figures

Figure 1. Map illustrating the known geographic distribution of Cyrtodactylus macrotuberculatus and C. phuketensis. Yellow star: the type locality of C. macrotuberculatus at Gunung Raya, Pulau Langkawi, Kedah, Malaysia. Green star: the type locality of C. phuketensis at Thalang District, Phuket Island, Phuket Province. Yellow circles: C. macrotuberculatus samples used in this study. Green circle: C. phuketensis samples used in this study. Yellow squares: the distribution of C. macrotuberculatus taken from Grismer et al. (2012), and Quah et al. (2019). The samples used correspond to those in Table 1.

Figure 4. Male holotype of Cyrtodactylus macrotuberculatus from Pulau Langkawi, Kedah, Peninsular Malaysia (ZRC 2.6754) in preservative A dorsal view B ventral view C tuberculate gular region and throat, and D enlarge femoroprecloacal scale row and pores.

Figure 5. Adult male Cyrtodactylus macrotuberculatus from Adang Island, Satun Province, Thailand (ZMKU R 00871) in life.

Figure 6. Adult male Cyrtodactylus macrotuberculatus from Adang Island, Satun Province, Thailand (ZMKU R 00871) in preservative A dorsal and B ventral views.

Figure 7. Variation in dorsal body band pattern of Cyrtodactylus macrotuberculatus from Thailand. From left to right, upper: ZMKU R 00878, ZMKU R 00873 from Adang Island, Satun Province; and ZMKU R 00887 from Rawi Island, Satun Province. Lower: ZMKU R 00889 from Rawi Island, Satun Province; ZMKU R 00896 and ZMKU R 00895 from Phuket Province.

Figure 8. Color in life of Cyrtodactylus macrotuberculatus from Thailand A adult male ZMKU R 00883 from Rawi Island, Satun Province B subadult female ZMKU R 00895 from Thalang District, Phuket Province, and C juvenile ZMKU R 00898 from Kathu District, Phuket Province.

Figure 9. Precloacal region in female specimens of Cyrtodactylus macrotuberculatus A paratype ZRC 2.6758, from Telaga Tujuh, Pulau Langkawi, Malaysia, and B ZMKU R 00896 from Thalang District, Phuket Province, Thailand.

Figure 10. Habitats of Cyrtodactylus macrotuberculatus in Thailand A Adang Island, Satun Province B Kathu District, Phuket Province, and C Hat Yai District, Songkhla Province.