Cyrtodactylus majulah, Grismer & Wood & Jr & Lim, 2012

Grismer, L. Lee, Wood, Perry L., Jr & Lim, Kelvin K. P., 2012, Cyrtodactylus Majulah, A New Species Of Bent-Toed Gecko (Reptilia: Squamata: Gekkonidae) From Singapore And The Riau Archipelago, Raffles Bulletin of Zoology 60 (2), pp. 487-499 : 490-496

Cyrtodactylus majulah , new species

( Figs. 3–5 )

Gymnodactylus marmoratus (not of Kuhl) – Sworder, 1925: 63; Smith, 1930: 12

Cyrtodactylus quadrivirgatus (not of Taylor) – Lim & Lim, 1992: 121; Baker & Lim, 2008: 80

Cyrtodactylus quadrivirgatus (in part) – Dring, 1979: 228; Denzer & Manthey, 1991: 314; Chan-ard et al., 1999: 24; Manthey & Grossman, 1997: 227; Teo & Rajathurai, 1997: 390; Cox et al., 1998: 89; Das, 2010: 214; Grismer, 2011a: 426; Grismer, 2011b: 118

Holotype. — Adult male, 67.9 mm SVL ( ZRC 2.6950 ); Singapore: Central Catchment Nature Reserve, Nee Soon Swamp-forest (1°22'48.99"N, 103°49'05.39"E; 51 m); coll: L. Grismer, E. Quah, K. O. Chan, A. Figueroa & A. Devan-Song, 22 Jun.2011.

Paratypes. — Adult male: 68.0 mm SVL ( ZRC 2.6951 ); adult male: 62.7 mm SVL ( ZRC 2.6952 ); adult male: 67.2 mm SVL ( LSUHC 10458 ); adult male: 67.2 mm SVL ( ZRC 2.6953 ); collection data as per holotype .

Non-types. — Singapore. 1 ex.: 50.9 mm SVL ( ZRC 2.1133 ); Botanic Gardens; coll: F. N. Chasen, Feb.1924 . 1 female: 59.0 mm SVL ( ZRC 2.2552 ); Nee Soon Swamp-forest; coll: K. Lim & P. K. L. Ng, 31 Dec.1989 . 1 ex.: 41 mm SVL ( ZRC 2.3190 ); Nee Soon Swamp-forest; coll: K. Lim, 14 Nov.1990 . 1 juvenile: 28.9 mm SVL ( ZRC 2.3205 ); Nee Soon Swamp-forest; coll: P. K. L. Ng, 23 Nov.1990 . 1 male: 68.0 mm SVL ( ZRC 2.3261 ); Nee Soon Swampforest; coll: P. K. L. Ng, 1 Jan.1991 . 2 ex.: 61.6– 40.3 mm SVL ( ZRC 2.3283 – 3284 ); forest at Upper Peirce Reservoir Park ; coll: R. C. H. Teo, 24 Oct.1991 . 1 female: 63 mm SVL ( ZRC 2.4576 ); Nee Soon Swamp-forest ; coll: T. M. Leong, 27 Feb.1999 . 1 female: 66.5 mm SVL ( ZRC 2.4862 ); forest at Upper Seletar Reservoir Park ; coll: T. M. Leong & C. P. Lim, 5 Jan.2001 . 1 ex.: 60 mm SVL ( ZRC 2.5643 ); forest at Upper Seletar Reservoir Park ; coll: T. M. Leong, 15 Mar.2003 . 1 male: 68.5 mm SVL ( ZRC 2.6796 ); Nee Soon Swamp-forest ; coll: T. M. Leong, T. O’Dempsey & S. H. Yeo, 2 Jan.2009 . Indonesia. 1 ex. ( ZRC 2.6555 ), 58 mm SVL; Riau Archipelago, Pulau Bintan , northern coastal forest; coll: T. M. Leong & R. Subaraj, 18 Oct.2007 .

Diagnosis. — Cyrtodactylus majulah new species is differentiated from its Sundaland congeners by having the unique combination of a maximum SVL of 68.0 mm; eight or nine supralabials; seven or eight infralabials; strongly tuberculated body and limbs; 39–46 paravertebral tubercles; 43–53 ventral scales; no abrupt contact of large and small postfemoral scales; no enlarged femoral scales; 20–23 subdigital lamellae; no femoral pores; no deep precloacal groove; enlarged precloacal scales; 7–11 precloacal pores; no enlarged median subcaudals; tubercles on the anterior portion of the tail; no reticulated pattern on head; having a blotched dorsal pattern; and lacking the paired, dark, semi-lunar shaped blotches on the upper nape prominently outlined in white. These and other characters are scored across all Sundaland species ( Table 4).

Description of holotype. — Adult male SVL 67.9 mm; head moderate in length (HL/SVL 0.27) and wide (HW/HL 0.64), flat (HD/HL 0.42), distinct from neck, and triangular in dorsal profile; lores weakly inflated, prefrontal region deeply concave, canthus rostralis rounded; snout elongate (ES/HL 0.44), rounded in dorsal profile; eye large (ED/HL 0.22); ear opening elliptical, obliquely oriented, moderate in size (EL/HL 0.09); eye to ear distance greater than diameter of eye; rostral square, partially divided dorsally by three small postrostral scales, bordered posteriorly by left and right supranasals, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by a large, anterior supranasal and smaller posterior supranasal, posteriorly by two postnasals and ventrally by first supralabial; eight (R,L) rectangular supralabials extending to just beyond upturn of labial margins tapering abruptly below midpoint of eye, first supralabial largest; eight (R,L) infralabials tapering smoothly posteriorly to beyond orbit; scales of rostrum and lores raised, slightly larger than granular scales on top of head and occiput; scales of occiput intermixed with slightly enlarged tubercles; dorsal superciliaries not elongate or keeled; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapazoidal postmentals which contact medially for 50% of their length posterior to mental; one row of slightly enlarged, elongate sublabials extending posteriorly to 6 th infralabial; gular scales small, granular, grading posteriorly into slightly larger, flatter, throat scales which grade into larger, flat, smooth, imbricate, pectoral and ventral scales.

Body relatively short (AG/SVL 0.45) with poorly defined ventrolateral folds; large postaxillary skin growth on right side; dorsal scales small, granular interspersed with moderately sized, conical, semi-regularly arranged, keeled tubercles; tubercles extend from occiput onto base of tail but no farther; tubercles on occiput and nape relatively small, those on body largest; approximately 22 longitudinal rows of tubercles at midbody between ventrolateral, body folds; 46 paravertebral tubercles on body; 43 flat, imbricate, ventral scales between ventrolateral, body folds, ventral scales larger than dorsal scales; precloacal scales large; no deep precloacal groove or depression.

lateralis

lateralis

Forelimbs moderate in stature, relatively short (FL/SVL 0.13); granular scales of forearm slightly larger than those on body, interspersed with slightly larger tubercles; palmar scales rounded, slightly raised; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae transversely expanded proximal to joint inflections, more granular distal to inflection; digits slightly more narrow distal to inflections; claws well-developed, claw base sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.17), covered dorsally by granular scales interspersed with large, conical tubercles and anteriorly by flat, slightly larger scales; ventral scales of thigh flat, imbricate, larger than dorsals; ventral tibial scales flat, imbricate; no rows of enlarged, flat, imbricate, femoral scales; small postfemoral scales do not form an abrupt union with large ventral scales of posteroventral margin of thigh; plantar scales low, flat; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae transversely expanded proximal inflections, more granular distal to inflections, digits more narrow distal to inflections; 21 subdigital lamellae on right 4 th toe, 22 on left; claws well-developed, base of claw sheathed by a dorsal and ventral scale.

Tail moderate, 88.1 mm in length, original, 6.5 mm in width at base, tapering to a point; dorsal scales of base of tail granular becoming flatter posteriorly; no median row of transversely enlarged subcaudal scales; subcaudal scales larger than dorsal caudal scales; one pair of paravertebral and dorsolateral tubercle rows on either side of midline on first one-fifth of tail; paravertebral tubercle rows not widely separated; caudal tubercles decrease in size posteriorly, extending approximately one-fifth length of tail; one enlarged, postcloacal tubercle at base of tail on hemipenal swelling; all postcloacal scales flat; partially everted hemipenes bilobed.

Colouration in life ( Fig. 4 ). — Dorsal ground colour of head, neck, trunk, limbs, and tail brown; straw-yellow mottling on head and rostrum extending from postorbital region onto nape as a poorly defined stripe; mottling coalesces on dorsum to isolate brown ground colour in four longitudinal rows of large, elongate blotches that extend to base of tail; mottling transitions into six yellowish caudal bands not encircling the the tail and separating five wider dark brown bands; posterior 30% of tail is uniformly dark brown; gular region, ventral surface of belly and limbs beige; each ventral scale bearing one or two small black spots; palmar and plantar regions grey; subcaudal region dark brown with irregularly shaped beige blotches; iris dark red.

Variation. — The general colouration and pattern of the paratypes closely match that of the holotype ( Fig. 5 ). ZRC 2.6952 tends to have a more transversely banded pattern as opposed to a longitudinally spotted pattern. ZRC 2.6953 has a regenerated tail that is uniformly light brown bearing a dense, darker stippling pattern. The posterior two-thirds of the tail of ZRC 2.6951 is missing. The light caudal bands of ZRC 2.6952 extend to the end of the tail whereas in the holotype (ZRC 2.6950) and LSUHC 10458, the bands extend only two-thirds of the length of the tail. The light mottling on the snout is slightly more distinct in ZRC 2.6555. Meristic differences are shown in Table 5.

Distribution. — Cyrtodactylus majulah new species is known from Singapore Island, only from the forested areas of the Central Catchment Nature Reserve, and on Pulau Bintan in the Riau Archipelago of Indonesia. Both islands are located off the southern end of the Malay Peninsula ( Fig. 1 ). ‘ Cyrtodactylus quadrivirgatus ’ reported from other parts of Singapore such as Bukit Timah Nature Reserve and Pulau Tekong (Teo & Rajathurai, 1997; Baker & Lim, 2008), apparently are not conspecific with C. majulah , and their identity will be re-evaluated elsewhere.

also been observed ( KKPL, pers. obs.) in other parts of the Central Catchment Nature Reserve in drier old secondary lowland forest. A specimen ( ZRC 2.1133 ) was obtained from the Botanic Gardens in 1924, but it seems to be extinct at that locality. ZRC 2.4576 is a gravid female carrying two eggs and was collected 27 Feb.1999 indicating that breeding occurs at least around this time of year .

Etymology. — In the Malay language (Bahasa Melayu), majulah means to go forward, to progress. This is in allusion to the present rapid advancement of research in the taxonomy of Cyrtodactylus , in which many species have recently been formally named, and more are waiting to be described.

Comparisons. — Cyrtodactylus majulah is distinguished from all its congeners on Sundaland except C. brevipalmatus (Smith) , C. cavernicolus Inger & King , C. elok Dring , C. ingeri Hikada , C. pubisulcus Inger , C. quadrivirgatus Taylor , and C. semenanjungensis Grismer & Leong , by having a maximum SVL of less than 70 mm. It differs from C. batucolus Grismer, Chan, Grismer, Wood & Belabut ; C. fumosus (Müller) ; C. leegrismeri Chan & Norhayati ; C. macrotuberculatus Grismer & Norhayati ; C. peguensis (Boulenger) ; C. pulchellus Gray ; C. quadrivirgatus ; and C. seribuatensis Youmans & Grismer , in having more than 42 ventral scales. From C. cavernicolus ; C. consobrinus (Peters) ; C. durio Grismer, Anuar, Quah, Muin, Chan, Grismer & Norhayati ; C. jarakensis Grismer, Chan, Grismer, Wood & Belabut ; C. lateralis (Werner) ; C. malayanus (de Rooij) ; and Natural history. — The type series was obtained in the Nee Soon Swamp Forest ( Corner, 1978), in the north-eastern part of the Central Catchment Nature Reserve that covers the central area of Singapore Island. All five specimens were found on the trunks of small trees in the vicinity of leafy cover 1.5–2 m above the ground. Four other specimens were seen in the same vicinity under the same conditions. One was seen 0.75 m above the ground on the surface of an exposed leaf. The habitat in which the type series was collected is a lowland equatorial rainforest criss-crossed by small streams ( Fig. 6 ). All specimens were found in dense vegetation in the vicinity of water courses. Cyrtodactylus majulah new species is not restricted to swamp-forest habitat, for it has C. stresemanni Rösler & Glaw ; it differs in having fewer than 52 ventral scales. From C. batucolus ; C. batik Iskandar, Rachmansah & Umilaela ; C. brevipalmatus (Smith) ; C. elok ; C. peguensis ; and C. yoshi Hikada ; it differs in having more than 19 subdigital lamellae on the fourth toe. It differs from C. jarakensis and C. yoshii in having fewer than 24 subdigital lamellae on the fourth toe. Cyrtodactylus majulah differs from all Sundaland Cyrtodactylus except C. aurensis Grismer , C. baluensis (Mocquard) , C. batik , C. consobrinus , C. ingeri , C. leegrismeri , C. macrotuberculatus , C. malayanus , C. peguensis , and C. pulchellus in not having enlarged subcaudal scales. It is further separated from C. aurensis , C. cavernicolus , C. durio , C. macrotuberculatus , C. marmoratus , C. pubisulcus , C. pulchellus , and C. stresemanni in not having a deep precloacal groove. Cyrtodactylus majulah has 7–11 precloacal pores which separates it from C. batucolus , C. batik , C. semenanjungensis , and C. seribuatensis Youmans & Grismer , which lack precloacal pores; and from C. lateralis , C. marmoratus , and C. durio which have more than 11.

Cyrtodactylus majulah new species had long been referred to as C. quadrivirgatus ( Lim & Lim, 1992; Teo & Rajathurai, 1997; Baker & Lim, 2008). Grismer (2011a) noted a significant departure in colour pattern between the Singapore population and that of C. quadrivirgatus from southern Peninsular Malaysia at Endau-Rompin, Johor and indicated that the conspecificity of these populations was under examination. Cyrtodactylus majulah new species differs from C. quadrivirgatus in having a dorsal pattern composed of large, semi-paired, paravertebral spots as opposed to stripes and broken bands; more ventral scales (43–51 vs. 34–42); lacking, as opposed to having enlarged femoral scales ( Fig. 7 ); and having 7–11 precloacal pores as opposed to 0–4 precloacal pores. Cyrtodactylus majulah new species is most similar to C. pantiensis from the nearby Gunung Panti Forest Reserve, Johor in southern Peninsular Malaysia but differs in having eight or nine supralabials as opposed to 10 or 11; having seven or eight infralabials as opposed to 9–11; 39–46 paravertebral tubercles as opposed to 35–37; having a maximum SVL of 68.0 mm as opposed to 77.2 mm; and lacking the paired, dark, semi-lunar shaped blotches on the upper nape prominently outlined in white that are diagnostic of C. pantienesis . There is also a sequence divergence of 17.8–18.1% between these two taxa.

Figures

Fig. 3. a) Cyrtodactylus semenanjungensis LSUDPC 4450 from Gunung Panti Forest Reserve, Johor, Malaysia. b) C. majulah LSUDPC 6307 from the Nee Soon Swamp, Singapore. c) C. pantiensis LSUDPC 4477 from Gunung Panti Forest Reserve, Johor, Malaysia. d) C. payacola LSUDPC 6267 from Bukit Panchor, Penang, Malaysia.

Fig. 4. Holotype of Cyrtodactylus majulah (ZRC 2.6950, male) from Central Catchment Nature Reserve, Nee Soon Swamp-forest, Singapore.

Fig. 5. Type series of Cyrtodactylus majulah (all males). From left to right: ZRC 2.6950 (holotype), ZRC 2.6951 (paratype), ZRC 2.6952 (paratype), LSUHC 10458 (paratype), and ZRC 2.6953 (paratype).

Fig. 1. Location of Cyrtodactylus majulah in Singapore and Pulau Bintin, Indonesia.

Fig. 6. Microhabitat of Cyrtodactylus majulah in the Nee Soon Swamp, Singapore.

Fig. 7. Cloacal region of Cyrtodactylus quadrivirgatus (LSUHC 8185) from Endau-Rompin, Johor, Malaysia, showing the large femoral scales (upper); and C. majulah (ZRC 2.6951) from the Central Catchment Nature Reserve, Nee Soon Swamp-forest, Singapore, showing the absence of enlarged femoral scales (lower).