Cyrtodactylus bintangrendah, Grismer, Lee, Wood, Perry L., Quah, Evan S. H., Anuar, Shahrul, Muin, Abdul, Sumontha, Montri, Ahmad, Norhayati & Bauer, Aaron M., 2012

Grismer, Lee, Wood, Perry L., Quah, Evan S. H., Anuar, Shahrul, Muin, Abdul, Sumontha, Montri, Ahmad, Norhayati & Bauer, Aaron M., 2012, A phylogeny and taxonomy of the Thai-Malay Peninsula Bent-toed Geckos of the Cyrtodactylus pulchellus complex (Squamata: Gekkonidae): combined morphological and molecular analyses with descriptions of seven new species, Zootaxa 3520, pp. 1-55 : 32-37

Cyrtodactylus bintangrendah sp. nov.

Bintang Lowland Bent-toed Gecko Figs. 13 , 14

Cyrtodactylus pulchellus Grismer 2011:416 (in part).

Holotype.—Adult male ( ZRC 2.6973) collected on 29 October 2011 by E. S. H. Quah, Fatim, S. B. M., Nor Amira, B. A. R. and Yusof O. from Bukit Mertajam, Seberang Perai, Penang, Peninsular Malaysia (05° 35.698 N, 100° 49.253 E) at 451 meters above sea level.

Paratypes.— Paratypes LSUHC 9974–76 (adult female, adult male, and adult female respectively) were collected from Lenggong, Perak. ZRC 2.6974 (adult male) was collected on 1 October 2010 by E. S. H. Quah from the type locality. Adult male ( LSUHC 10520) bears the same collection data as the holotype. Juvenile female ( ZRC 2.6975) collected on 20 June 2010 by E. S. H. Quah from Bukit Palong, Kedah, Peninsular Malaysia (5° 36.140 N 100° 55.190 E) at 238 meters in elevation. Adult male ( LSUHC 10331) collected on 26 July 2011 by E. S. H. Quah at Ulu Paip, Kedah (5° 24.630 N 100° 38.419 E) at 200 meters above sea level as estimated from Google Earth.

Diagnosis.—Adult males reaching 114.4 mm SVL, adult females reaching 103.1 mm SVL; 8–12 supralabials, 8–10 infralabials; tubercles of dorsum small to moderate with no intervening smaller tubercles; no tubercles on ventral surfaces of forelimbs, gular region, or in ventrolateral body fold; 33–44 paravertebral tubercles; 22–25 longitudinal rows of dorsal tubercles; 31–39 rows of ventral scales; 20–24 subdigital lamellae on fourth toe; 41–46 femoro-precloacal pores in males; dorsum not bearing a scattered pattern of white tubercles; four, rarely five, body bands in adults lacking lightened centers and light colored tubercles; band to interspace ratio 0.50–1.25; 8–14 dark caudal bands on original tail; light caudal bands in adults not heavily infused with dark pigment; and posterior portion of tail in hatchlings and juveniles banded not white. These characters are scored across all species of the Cyrtodactylus pulchellus complex in Table 6 .

Description of holotype.—Adult male SVL 114.4 mm; head large, moderate in length (HL/SVL 0.27) and wide (HW/HL 0.81), somewhat flattened (HD/HL 0.49), distinct from neck, and triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; frontal and prefrontal regions deeply concave; canthus rostralis rounded anteriorly; snout elongate (ES/HL 0.45), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.24); ear opening elliptical, moderate in size (EL/HL 0.05), vertically oriented; eye to ear distance greater than diameter of eye; rostral rectangular, divided dorsally by a large postrostral (=internasal), bordered posteriorly by left and right supranasals, a medial postrostral, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by a large anterior supranasal and small posterior supranasal, posteriorly by two postnasals, ventrally by first supralabial; 9,10(R,L) square supralabials extending to just beyond upturn of labial margin, tapering abruptly below midpoint of eye; first supralabial largest; 8(R,L) infralabials 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 of occiput and top of head posterior to eyes intermixed with small 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, rectangular postmentals which contact medially for 30% of their length; two rows of enlarged, elongate sublabials extending posteriorly to 6th infralabial; small, granular, gular scales grading posteriorly into larger, flat, smooth, imbricate, pectoral and ventral scales.

Body relatively short (AG/SVL 0.44) with well-defined, weakly tuberculate, ventrolateral folds; dorsal scales small, granular, interspersed with large, trihedral, regularly arranged, keeled tubercles; smaller intervening tubercles generally absent; tubercles extend from top of head posterior to eyes to caudal constriction and onto tail where they occur in transverse rows separated by 6–8 small, flat scales; caudal tubercles largest dorsally, absent laterally and ventrally; caudal tubercles decrease in size posteriorly; tubercles on occiput and nape relatively small, those on body largest; approximately 24 longitudinal rows of tubercles at midbody; 39 paravertebral tubercles; 36 flat imbricate ventral scales between ventrolateral body folds, ventral scales larger than dorsal scales; precloacal scales large, smooth; distinct precloacal groove.

Forelimbs moderate, relatively short (FL/SVL 0.16); scales on dorsal surfaces of forelimbs raised, intermixed with larger tubercles; scales of ventral surface of forearm flat, subimbricate, lacking tubercles; palmar scales rounded; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae rectangular proximal to joint inflection, only slightly expanded distal to inflection; digits more narrow distal to joints; claws welldeveloped, sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.20), larger tubercles on dorsal surface of thigh separated by smaller granular scales, tubercles on dorsal surfaces of foreleg same size as those on thigh; ventral scales of thigh flat, smooth, imbricate, larger than dorsal granular scales; ventral, tibial scales flat, smooth, imbricate; single row of greatly enlarged, flat, rectangular, imbricate, pore-bearing femoral scales extend from knee to knee through precloacal region where they are continuous with enlarged, pore-bearing precloacal scales; 42 contiguous, pore-bearing femoro-precloacal scales forming an inverted T bearing a deep, precloacal groove in which 13 pore-bearing scales are found (six on right, seven on left side of groove); postfemoral scales immediately posterior to pore-bearing scale row small, forming an abrupt union with pore-bearing, postfemoral scales on posteroventral margin of thigh; plantar scales low, slightly raised; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae proximal to joint inflection rectangular, only slightly expanded distal to inflection; digits more narrow distal to joints; claws welldeveloped, sheathed by a dorsal and ventral scale; 23(R) 24(L) subdigital lamellae on 4th toe.

Tail 140 mm in length, 10.1 mm in width at base, tapering to a point; dorsal scales of tail flat, squarish; tail segmented with 6–8 transverse scale rows per segment; posterior margin of segments bordered by 4–6 larger tubercles dorsally in anterior one-third of tail, fewer posteriorly; subcaudal region bearing large, transverse scales; shallow dorsal and lateral caudal furrows extend entire length of tail; base of tail bearing hemipenial swellings; 3,2(R,L) small, postcloacal tubercles on hemipenial swellings; postcloacal scales smooth, flat, large, imbricate.

Coloration in life. Dorsal ground color of head, body, limbs, and tail tan; no white chevron on rostrum; wide, brown, nuchal band extends from posterior margin of one eye to posterior margin of other eye, edged with prominent thin, yellow, lines; four similarly colored dorsal bands between limb insertions with immaculate, yellow, edging; first band terminates at shoulders, second and third bands terminate just dorsal to ventrolateral fold, fourth band terminates on anterior margin of hind limb insertions; body band/interspace ratio 1.25; one additional dark brown band posterior to hind limbs; no band on posterior margin of thigh; four wide, dark bands approximately three times the width of four white bands on original potion of tail; all bands encircle tail; regenerated portion of tail dark brown, immaculate; no bands on tail or body have lighten centers or encompass prominent yellowish to white tubercles; ventral surfaces of head, limbs, and tail beige; gular scales stippled; abdomen immaculate, except for slightly darker, lateral regions, and weakly stippled ventral scales.

Variation. ZRC 2.6975 (subadult female; SVL 67 mm from Bukit Palong, Kedah) closely approaches the holotype in general aspects of coloration except that its dorsal pattern is more contrasting being that the specimen is not an adult ( Fig. 14 ). The paratypes from Lenggong, Perak (LSUHC 9974–76) have an overall orangish brown dorsal coloration like the holotype but are less contrasting in that the light edging of the bands is less distinct, and their dorsal tubercles are lower in profile and less distinct. Additionally, LSUHC 9974 and 9976 have five body bands and LSUHC 9975 has 14 dark and 13 whitish caudal bands, respectively. Meristic differences in the type series and additional specimens examined are presented in Table 11.

Additional specimen examined. —One additional specimen (LSUHC 9977) is a subadult female (SVL 74 mm) from Lenggong which is also lighter overall but has a more contrasting color pattern with more distinctive band edging with respect to the adults from Lenggong owing to its smaller size ( Fig. 14 ).

Distribution. Cyrtodactylus bintangrendah sp. nov. is presumed to range throughout lowland forests up to 450 meters in elevation through the foothills and lower slopes surrounding the Bintang Mountain Range. Along the west side of the mountains it ranges from at least Bukit Mertajam, Seberang Perai, Penang in the south to Bukit Palong, Kedah in the north ( Fig. 3 ). East of the Bintang range, it is known only from the Lenggong Valley, Perak ( Fig. 3 ). Grismer (2011:424) reported lowland populations of C. pulchellus sensu strictu from much farther south in Selangor, Negeri Sembilan, and Johor which may indicate that C. bintangrendah sp. nov. shares a similar distribution pattern proposed for other lowland species found west of the Banjaran Titiwangsa ( Grismer 2011).

Natural history. Cyrtodactylus bintangrendah sp. nov. is a nocturnal species found in lowland and lower hill dipterocarp forest below 500 meters elevation. At Bukit Mertajam, Penang, specimens were encountered on large rocks and boulders especially along forest streams at night ( Fig. 13 ). Lizards were also observed on tree trunks and vines close to boulders but escaped into crevices in the rocks when approached. ZRC 2.6974 was found crawling on a retaining rock wall along a road cut at night at Bukit Mertajam. At Bukit Panchor, Penang, a male and a female (LSUDPC 6320) were collected at night crawling over large granite boulders that formed cave-like environments in lowland forest. Farther north at Bukit Palong, Kedah, the single subadult female (ZRC 2.6975) was found in lower hill dipterocarp forest at night clinging onto a sapling about 30cm above the ground. Other specimens were observed crawling on rocks and tree trunks. The male from Ulu Paip, Kedah (LSUHC 10331), was found on the side of a tree trunk at night approximately 1m off the ground near a river edged with boulders. Another specimen was observed in the vicinity on another tree trunk but ran into a crevice at the base of the trunk when approached. East of the Bintang Range at Lenggong Valley, Perak, C. bintangrendah sp. nov. is saxicolous and found on limestone karst outcrops and around the mouths of limestone caves. During the day geckos are occasionally observed hiding in cracks and crevices of the outcrops. At night, lizards emerge and can be found perched or crawling over the walls of the cave 1–3m above the ground.

Etymology. The word bintangrendah is a combination of the Malaysian words bintang meaning star and rendah meaning low and is in reference to the lowland regions surrounding the Bintang Mountain Range wherein this species occurs; bintangrendah is a noun in apposition and is thus invariable.

State, Malaysia. m=male; f=female; SVL=snout-vent length; TL=tail length; TW=tail width; FL=forelimb length;

TBL=tibia length; AG=axilla-groin length; HL=head length; HW=head width; HD=head depth; ED=eye diameter;

EE=eye to ear distance; ES=eye to snout distance; EN=eye to nostril distance; IO=interorbital distance; EL=ear length;

and IN=internarial distance; w=weak; /=data unavailable.

LSUHC LSUHC LSUHC ZRC ZRC ZRC LSUHC LSUHC LSUHC ZRC 9974 9975 9976 2.6974 2.6975 2.6973 10520 10331 9977 2.1177 Comparisons. Cyrtodactylus bintangrendah sp. nov. is separated from C. macrotuberculatus in lacking large tubercles on the dorsal surface of the head, body and limbs, on the underside of the forearms, in the gular region, and having 31–39 ventral scales as opposed to 17–28 ventral scales. From C. astrum sp. nov., C. langkawiensis sp. nov., C. lekaguli sp. nov. and C. pulchellus , C. bintangrendah sp. nov. differs in having more than 40 femoroprecloacal pores. It differs from C. trilatofasciatus sp. nov. in having four or five body bands with a band/ interspace ratio of 0.50–1.25 as opposed to having three bands with a band/interspace ratio of 2.00–2.75. Cyrtodactylus bintangrendah sp. nov. is further separated from C. astrum sp. nov. in lacking a scattered dorsal pattern of white tubercles. Hatchling and juvenile C. bintangrendah sp. nov. do not have a white tail tip which separates it from C. astrum sp. nov., C. langkawiensis sp. nov. and C. lekaguli sp. nov. whose hatchlings and juveniles do. Cyrtodactylus bintangrendah sp. nov. is separated from C. trilatofasciatus sp. nov. by having 8–14 dark caudal bands vs. six or seven in C. trilatofasciatus sp. nov. Its larger maximum SVL of 114.4 mm differentiates it further from C. langkawiensis sp. nov. and C. lekaguli sp. nov. which do not reach over 104 mm SVL and from the much larger C. trilatofasciatus sp. nov. (maximum SVL 122.2 mm). Cyrtodactylus bintangrendah sp. nov. cannot be discretely separated from C. bintangtinggi sp. nov. based on morphology. However, student t- test indicates the mean number of femoro-precloacal pores in C. bintangrendah sp. nov. (42.8) is significantly greater (p <0.003) than that of C. bintangtinggi sp. nov. (39.7). Additionally the average sequence divergence between these two species is 6.9–8.0%.

Remarks. Along the west side of the Banjaran Bintang, Cyrtodactylus bintangrendah sp. nov. is associated with granite outcroppings in lowland forest. However, the population in the Lenggong Valley, Perak occurs along the eastern side of the Banjaran Bintang in karst formations and the color pattern of the lizards from this area has converged to some extent on that of the other karst dwelling species, C. astrum sp. nov., C. langkawiensis sp. nov. and C. lekaguli sp. nov. in being generally lighter overall and have a significantly higher number of dark caudal bands.

Figures

FIGURE 3. Distribution of the species of the Cyrtodactylus pulchellus complex. Closed circles represent localities from which specimens have been examined genetically and / or morphologically (see appendix and Table 1). Closed squares are localities from which specimens have been only examined morphologically or photographically (see appendix). Closed stars represent the type localities from where specimens have been examined genetically, morphologically and photographically. 1. Suwankuha Cave, Muang District, Phang-nga Province, Thailand. 2. Petch Phanomwung Cave, Kanchanadit District, Surat Thani Province, Thailand. 3. Phuket Island 4. Khao Chong waterfall, Khao Chamao, Khaochong District, Trang Province, Thailand. 5. Phattalung Province, Thailand. 6. Rattabhumi District, Songkhla, Thailand. 7. La-ngu District, Satun Province, Thailand (three squares). 8. Gua Wang Burma, Perlis State Park, Perlis, Malaysia (two circles). 9. Wang Kelian, Perlis, Malaysia. 10. Adang Island, Satun Province, Thailand. 11. Pulau Langkawi, Kedah, Malaysia (two stars). 12. Gua Kelam and Bukit Chabang, Malaysia (two circles). 13. Kuala Perlis, Perlis, Malaysia. 14. Chuping, Perlis, Malaysia. 15. Bukit Wang, Kedah, Malaysia. 16. Ulu Muda, Kedah, Malaysia. 17. Gunung Jerai and Hutan Lipur Sungai Tupah, Kedah, Malaysia (two circles). 18. Bukit Palong, Perak, Malaysia. 19. Ulu Paip, Kedah, Malaysia. 20. Bukit Mertajam, Penang, Malaysia. 21. Pulau Pinang, Penang, Malaysia. 22. Lengong Valley, Perak, Malaysia. 23. Bukit Panchor, Penang, Malaysia. 24. Bukit Larut, Perak, Malaysia. 25. Bukit Pondok, Perak, Malaysia. 26. Ringlet, Cameron Highlands, Pahang, Malaysia. 27. Gunung Bubu, Perak, Malaysia. 28. Fraser’s Hill, Pahang, Malaysia. 29. Genting Highlands, Pahang, Malaysia. 30. Gunung Angsi, Negeri Sembilan, Malaysia. P = Tham Phung Chang Cave, Phang-nga City, Muang District, Phang-nga Province, Thailand. U = Nakon Si Thammarat Province; specific locality unknown.

FIGURE 13. Cyrtodactylus bintangrendah. Upper left: ZRC 2.6974 (paratype); adult male from Bukit Mertajam, Penang, Malaysia (photo by E. Quah). Upper right: LSUDPC 6320; adult female from Bukit Panchor, Penang, Malaysia (photo by E. Quah). Middle left: LSUHC 9974 adult male (paratype) from Lenggong Valley, Perak, Peninsular Malaysia (photo by M. Muin). Middle right: LSUHC 9977; juvenile from Bukit Mertajam, Penang, Malaysia (photo by E. Quah). Lower left: granite habitat of C. bintangrendah at Bukit Mertajam, Penang, Malaysia (photo by E. Quah). Lower right: karst habitat of C. bintangrendah at Gua Puteri, Lenggong Valley, Perak, Malaysia (photo by M. A. Muin).

FIGURE 14. Type series and additional specimens examined of Cyrtodactylus bintangrendah. Upper row from left to right: holotype ZRC 2.6973 and paratypes LSUHC 10520 and ZRC 2.6974 from Bukit Mertajam; paratypes ZRC 2.6975 from Palong, LSUHC 10331 from Ulu Paip, ZRC 2.117. Additional specimen examined; ZRC 2.1177 from Gunung Pondok. Lower row: paratypes LSUHC 9974, 9975, 9976 from Lenggong Valley. Additional specimen examined; LSUHC 9977 from Lenggong Valley.