Cyrtodactylus macrotuberculatus, Grismer, Lee & Ahmad, Norhayati, 2008

Grismer, Lee & Ahmad, Norhayati, 2008, A new insular species of Cyrtodactylus (Squamata: Gekkonidae) from the Langkawi Archipelago, Kedah, Peninsular Malaysia, Zootaxa 1924, pp. 53-68 : 55-64

Cyrtodactylus macrotuberculatus sp. nov.

Figures 2–7

Holotype. Adult male ( ZRC 2.6754) collected by L. Lee Grismer, Perry L. Wood, Jr., Timothy A. Youmans, and Norsham Yaakob on 8 August 2005 from beneath an overpass on the road to Gunung Raya (06° 23.023 N, 99° 49.126 E; 621 m asl.), Pulau Langkawi, Kedah, Peninsular Malaysia.

Paratypes. All paratypes were collected by the same collectors. ZRC 2.6757 was collected on 9 August 2005 from Telaga Tujuh (06° 22.522 N, 99° 40.236 E; 100 m asl.); ZRC 2.6755 and 2.6756 have the same collection data as the holotype; and ZRC 2.6758 was collected at Lubuk Semilang (06° 20.551 N, 99° 48.342 E; 61 m asl.) on 21 August 2005.

Diagnosis. Cyrtodactylus macrotuberculatus is distinguished from all other Sunda Shelf species by having a maximum SVL of 120 mm; very large, trihedral, keeled, tubercles on body, limbs, and tail; tubercles present on occiput and top of head; no granular scales (only large tubercles) on dorsal surfaces of limbs; 19– 22 ventral scales; transversely enlarged, median, subcaudal scales; proximal subdigital lamellae square; 21–23 subdigital lamellae on fourth toe; abrupt transition between posterior and ventral femoral scales; enlarged femoral and precloacal scales with a continuous series of 35–37 pore-bearing scales (femoero-precloacal scales); precloacal groove present, precloacal depression absent; no white recticulum on head; body and tail banded, limbs immaculate. The majority of these characters are summarized across all Sunda Shelf species in Table 1 . The others are discussed below (see Comparisons).

Description of holotype ( Fig. 2 ). Adult male SVL 119.0 mm; head large, moderate in length (HL/SVL 0.28) and wide (HW/HL 0.72), somewhat flattened (HD/HL 0.43), distinct from neck, and triangular in dorsal profile; lores concave; frontal and prefrontal regions deeply concave; canthus rostralis sharply rounded; snout elongate (ES/HL 0.42), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.23); ear opening elliptical, moderate in size (EL/HL 0.08), obliquely oriented; eye to ear distance greater than diameter of eye; rostral square, deeply divided dorsally, bordered posteriorly by left and right supranasals and medial postrostral (=internasal), bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by a large, anterior supranasal and small, posterior supranasal, posteriorly by granular scales, ventrally by first supralabial; 12(R) 11(L) square supralabials extending to just beyond upturn of labial margins tapering abruptly below midpoint of eye; first supralabial largest; eight (R,L) infralabials not tapering in size anterior to orbit; scales of rostrum and lores raised, larger than granular scales on top of head and occiput, those on canthus rostralis largest; scales of occiput intermixed with enlarged tubercles; large, boney frontal ridges bordering orbit confluent with boney, transverse, parietal ridge; dorsal superciliaries elongate, smooth, V-shaped; mental triangular, bordered laterally by first infralabials and posteriorly by left and right, rectangular postmentals which contact medially for 50% of their length; one row of slightly enlarged, elongate sublabials extending posteriorly to 5th 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) with well-defined, tuberculate, ventrolateral folds; dorsal scales small, granular, interspersed with very large, trihedral, regularly arranged, keeled tubercles separated by no more than three granules at their base; tubercles extend from occiput to caudal constriction but not onto regenerated tail; tubercles on occiput and nape relatively small, those on body largest; approximately 23 longitudinal rows of tubercles at midbody; 45 paravertebral tubercles; 21 flat, imbricate, large, ventral scales between ventrolateral, body folds, ventral scales much larger than dorsal scales; precloacal scales large, smooth; distinct precloacal groove ( Fig. 5 ).

Forelimbs moderately robust, relatively short (FL/SVL 0.16); virtually no granular scales on dorsal surfaces of forelimbs, only large, trihedral, keeled tubercles; palmar scales 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 a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.17), virtually no granular scales on dorsal surfaces of hind limbs, only large, trihedral, keeled tubercles; ventral scales of thigh flat, imbricate, same size as dorsals; ventral, tibial scales flat, imbricate, slightly keeled; two rows of enlarged, flat, imbricate, femoral scales extend from knee to knee through precloacal region where they are continuous with enlarged, precloacal scales; posterior row of enlarged femoral scales composed of 36 contiguous, pore-bearing scales (femoero-precloacal pores) extending from knee to knee forming a V bordering the deep, precloacal groove; nine pores bordering groove; postfemoral scales immediately posterior to the porebearing 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; 21 (R,L) subdigital lamellae on 4th toe.

tubercles on head and/or 1 1 1 1 1 1 1 occiput

tubercles on at least 1/3 of tail 1 — 1 0 1 1 1 Tail regenerated, 96 mm in length, 11.9 mm in width at base, tapering to a point; dorsal scales at base of tail square, smooth, flat, subimbricate, lacking tubercles; median row of transversely enlarged, subcaudal scales; three small, postcloacal tubercles at base of tail on hemipenial swellings; all postcloacal scales flat, large, imbricate.

Coloration in life. Dorsal ground color of head, body, limbs, and tail tan; wide, dark brown, nuchal band edged with thin, white lines extends from posterior margin of one eye to posterior margin of other eye; four similar dorsal bands between limb insertions, first band terminates at shoulders, second and third bands terminate just dorsal to ventrolateral fold, fourth band terminates in groin; one additional dark brown band posterior to hind limbs; no bands on regenerated tail; ventral surfaces of head, limbs, and tail smudged with brown; abdomen immaculate, beige except for slightly darker, lateral regions.

Variation. The paratypes match the holotype in all aspects of dorsal banding except they have nine or 10 dark bands on their original tails ( Fig. 3 ). The ground color of the body of the juvenile ZRC 2.6757 (SVL 51.2 mm) is bright yellow and all its bands are black with yellow edging that is difficult to discern ( Fig. 3 ). As this species ages, the ground color darkens to tan and the yellow edging on the bands turns white as seen in the subadult male ZRC 2.6756 (SVL 99.2 mm; Fig. 3 ). This subadult coloration transforms into the adult color pattern seen in the adult female ZRC 2.6755 (SVL 120 mm) with a tan ground color and dark brown, dorsal bands ( Fig. 4 ). All paratypes had complete tails with two rows of dorsal and lateral caudal tubercles (four rows in total) extending nearly the entire length of the tail. Although the degree (i.e., relative size) of tuberculation of the juvenile (ZRC 2.6757) is not nearly as extreme as that in the adults, the distribution of tubercles on the body is the same. Females lack a precloacal groove. Meristic differences in the type series are presented in Table 2 .

Distribution. Cyrtodactylus macrotuberculatus is known only from Pulau Langkawi in the Langkawi Archipelago, Kedah, Peninsular Malaysia ( Fig. 1 ) and is presumably endemic to this archipelago. This species is expected to occur on some of Pulau Langkawi’s smaller, satellite islands.

Natural history. Cyrtodactylus macrotuberculatus is a common, scansorial species found in nearly all forested habitats ranging from sea level to the summit of Gunung Machinchang at 700 m. Specimens have been observed abroad only at night and have been found in karst formations taking refuge in the rock cracks on Gunung Machinchang, on granite boulders along water courses at Lubuk Semilang and Telaga Tujuh, crossing the road leading to the summit of Gunung Raya, beneath overpasses and on road cuts along this road, and on road cuts and three meters above the ground on trees at Lubuk Semilang.

Etymology. The specific epithet macrotuberculatus comes from the Greek makros meaning long or large and the Latin tuberculum meaning a protuberance or knob and refers to the numerous, large tubercles covering this species body.

Comparisons. Cyrtodactylus macrotuberculatus is easily distinguished from all other Sunda Shelf species of Cyrtodactylus by having large tubercles in the gular region and throat as opposed to only granular scales ( Fig. 5 ) and by having relatively fewer but larger ventral scales (19–22 vs 29–65 for all other species collectively; Table 1 ). Although C. macrotuberculatus has masqueraded under the nomen of C. pulchellus ( Chan-ard et al. 1999; Manthey & Grossmann 1997; Grismer 2008b; Grismer et al. 2006a; Zimmerer 2004), it is clearly differentiable from that species by its much larger and more widely distributed tubercles ( Figs. 4–7 ); having a maximum SVL length of 120 mm vs 115 mm; having fewer ventral scales (19–22 vs 33–55); having square, proximal subdigital lamellae as opposed to rectangular lamellae ( Fig. 7 ); having more subdigital lamellae (21–23 vs 19–20); having large tubercles in the gular and throat regions as opposed to only having granular scales in those regions ( Fig. 5 ); having the ventrolateral body folds being composed of large, conical, tuberculate scales as opposed to being composed of small, smooth scales ( Fig. 7 ); having only large tubercles on the dorsal portions of the limbs as opposed to smaller tubercles and granular scales ( Fig. 6 ); having tubercles on the lateral portions of the abdomen as opposed to having only smooth, flat imbricate scales ( Fig. 7 ); and having much larger, postcloacal scales ( Fig. 5 ).

Figures

FIGURE 1. Distribution of Cyrtodactylus macrotuberculatus on Pulau Langkawi in the Langkawi Archipelago, Kedah, Peninsular Malaysia.

FIGURE 2. Dorsal view of holotype of Cyrtodactylus macrotuberculatus (ZRC 2.6754).

FIGURE 3. Upper: Juvenile female paratytpe of Cyrtodactylus macrotuberculatus (ZRC 2.6757, SVL 51.2 mm) showing the yellow ground color and black dorsal bands edged in yellow. Lower: subadult male paratytpe of Cyrtodactylus macrotuberculatus (ZRC 2.6756, SVL 99.2 mm) showing the tan ground color and dark brown dorsal bands edged in yellow.

FIGURE 4. Upper: Adult female paratytpe of Cyrtodactylus macrotuberculatus (ZRC 2.6755, SVL 120.0 mm) showing the light brown ground color and brown dorsal bands edged in white as well as its strong tuberculation. Lower: Adult male Cyrtodactylus pulchellus (LSUHC 8090, SVL 113.1 mm) from Fraser’s Hill, Pahang showing its overall weak tuberculation.

FIGURE 5. Upper left: holotytpe of Cyrtodactylus macrotuberculatus (ZRC 2.6754) showing tuberculate gular region and throat. Upper right: Cyrtodactylus pulchellus (LSUHC 8089) from Fraser’s Hill, Pahang showing smooth gular region. Middle: holotytpe of Cyrtodactylus macrotuberculatus (ZRC 2.6754) showing large femoero-precloacal scales and large postcloacal scales. Lower: Cyrtodactylus pulchellus (LSUHC 8089) from Fraser’s Hill, Pahang showing small femoero-precloacal scales and small postcloacal scales.

FIGURE 6. Upper: holotytpe of Cyrtodactylus macrotuberculatus (ZRC 2.6754) showing strongly tuberculate dorsum. Middle: Cyrtodactylus pulchellus (LSUHC 8089) from Fraser’s Hill, Pahang showing weakly tuberculate dorsum. Lower left: holotytpe of Cyrtodactylus macrotuberculatus (ZRC 2.6754) showing strongly tuberculate brachial region and absence of granular scales. Lower right: Cyrtodactylus pulchellus (LSUHC 8089) from Fraser’s Hill, Pahang showing weakly tuberculate brachial region with granular scales.

FIGURE 7. Upper: holotytpe of Cyrtodactylus macrotuberculatus (ZRC 2.6754) showing large, conical, tuberculate scales of ventrolateral fold and tuberculate scales on lateral margins on abdomen. Middle: Cyrtodactylus pulchellus (LSUHC 8089) from Fraser’s Hill, Pahang showing small, smooth scales of ventrolateral fold and small, smooth, flat, imbricate scales of lateral margins of abdomen. Lower left: holotytpe of Cyrtodactylus macrotuberculatus (ZRC 2.6754) showing the more oval shape of the proximal subdigital lamellae of the left foot. Lower right: Cyrtodactylus pulchellus (LSUHC 8089) from Fraser’s Hill, Pahang showing showing the more rectangular shape of the proximal subdigital lamellae of the left foot.

Tables

TABLE 1. Mensural and meristic data from Sunda Shelf species of Cyrtodactylus from peninsular Malaysia, Borneo, Java, and Sumatra. 1 = presence of character state, 0 = absence of character state, — data unavailable.

aurensis	baluensis	brevipalmatus	cavernicolous	consobrinus	elok	fumosus
SVL 92–95	72–86	64–73	64–81	97–121	79–82	71–75
tuberculation moderate to 0 strong	1	1	1	1	0	1
tubercles on forelimbs 0	1	1	0	1	1	1
tubercles on hind limbs 0	1	1	1	1	1	1
tubercles on head and/or 0 occiput	1	0	1	1	0	1
tubercles on at least 1/3 of tail 0	0	1	1	1	1	0
ventral scales 45–51	40–45	35–45	51–58	58–65	44–49	35–40
enlarged median subcaudals 1	1	0	0	1	0	0
proximal subdigital lamellae 1 broad	1	1	1	1	1	1
subdigital lamellae on 4th toe 18–23	21–23	16–19	22–26	23–28	18–23	20–22
contact of posterior thigh 0 scales abrupt	1	1	0	0	0	0
enlaged femoral scales 0	1	1	0	1	0	1
femoral pores 0	6–9	6,7	0	1–6	0	42–52
precloacal groove 1	0	0	1	0	0	0
enlarged precloacal scales 1	1	1	1	1	1	1
precloacal pores 7	9,10	9,10	4	9,10	8	42–52
precloacal and femoral pores/ 0 scales continuous	0	0	0	0	0	1
reticulate pattern on head 1	1	0	1	1	0	0
body banded 1	0	0	1	1	0	0
body blotched 0	1	1	0	0	1	1
body striped 0	0	0	0	0	0	0
contined.