Cyrtodactylus langkawiensis, 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 : 23-27

Cyrtodactylus langkawiensis sp. nov.

Langkawi Island Bent-toed Gecko Figs. 9 , 10

Holotype.—Adult male ( ZRC 2.6966) collected on 11 October 2008 by L. L. Grismer, Norhayati A., Chan, K. O., D. Belabut, and J. L. Grismer from Wat Wanaram, Pulau Langkawi, Kedah, Peninsular Malaysia (06° 20.275 N, 99° 52.507 E) at 35 meters above sea level.

Paratypes.—Adult females ( ZRC LSUHC 2.6967–68 and LSUHC 9123–24) collected from same locality on the same date as the holotype and by the same collectors; adult female ( ZRC 2.6969) collected at same locality as holotype by Chan, K. O., C. Johnson, and L. Grismer on 19 September 2009.

Diagnosis.—Adult males reaching 93.3 mm SVL, adult females reaching 99.8 mm SVL; 9–12 supralabials, 8–10 infralabials; tubercles of dorsum small to moderately large with no intervening smaller tubercles; no tubercles on ventral surfaces of forelimbs, gular region, or in ventrolateral body fold; 34–44 paravertebral tubercles; 21–25 longitudinal rows of dorsal tubercles; 38–43 rows of ventral scales; 19–21 subdigital lamellae on fourth toe; 30 femoro-precloacal pores in males; dorsum not bearing a scattered pattern of white tubercles; four rarely five dark body bands in adults lacking lightened centers and light colored tubercles; band to interspace ratio 0.75–1.00; 11–16 dark caudal bands on original tail; white caudal bands in adults infused with dark pigmentation; and posterior portion of tail in hatchlings and juveniles white. These characters are scored across all species of the Cyrtodactylus pulchellus complex in Table 6 .

Description of holotype.—Adult male SVL 93.3 mm; head large, moderate in length (HL/SVL 0.27) and wide (HW/HL 0.70), somewhat flattened (HD/HL 0.47), distinct from neck, and triangular in dorsal profile; lores flat anteriorly, inflated posteriorly; frontal and prefrontal regions concave; canthus rostralis rounded anteriorly; snout elongate (ES/HL 0.50), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.28); ear opening elliptical, moderate in size (EL/HL 0.11), vertically oriented; eye to ear distance greater than diameter of eye; rostral rectangular, divided dorsally by an inverted Y-shaped furrow, bordered posteriorly by left and right supranasals, a medial postrostral (=internasal), and an anomalous series of granular scales extending into dorsal portion of rostral; rostral bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by a large anterior supranasal and small posterior supranasal, posteriorly by single postnasal, ventrally by first supralabial; 12(R) 11(L) square supralabials extending to just beyond upturn of labial margin, tapering abruptly below midpoint of eye; second supralabial slightly larger than first; 10(R) 9(L) infralabials tapering in size posteriorly, second and third left infralabials fused; scales of rostrum and lores flat to slightly raised, larger than granular scales on top of head and occiput, those on posterior portion of canthus rostralis slightly larger; scales of occiput intermixed with small tubercles; large, boney frontal ridges bordering orbit confluent with boney, Vshaped, 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 75% of their length; one row of slightly enlarged, elongate sublabials extending posteriorly to 7th infralabial; small, granular, gular scales grading posteriorly into larger, flat, smooth, imbricate, pectoral and ventral scales.

Body relatively short (AG/SVL 0.46) with well-defined, non-tuberculate, ventrolateral folds; dorsal scales small, granular, interspersed with large, trihedral, regularly arranged, keeled tubercles, smaller intervening tubercles absent; tubercles extend from occiput to caudal constriction and onto tail where they occur in transverse rows separated by 6–8 small, flat scales; caudal tubercles largest dorsally, weak laterally, and absent ventrally; caudal tubercles decrease in size posteriorly, absent from regenerated portion of tail; tubercles on occiput and nape relatively small, those on body largest; approximately 23 longitudinal rows of tubercles at midbody; 34 paravertebral tubercles; 41 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 flat to granular, intermixed with a few larger tubercles; scales of ventral surface of forearm flat, rounded, 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 well-developed, 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 smaller; 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, porebearing femoral scales extend from knee to knee through precloacal region where they are continuous with enlarged, pore-bearing precloacal scales; 30 contiguous, pore-bearing femoro-precloacal scales forming an inverted T bearing a deep, precloacal groove in which five pore-bearing scales are found (three on right, two 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 well-developed, sheathed by a dorsal and ventral scale; 20 (R,L) subdigital lamellae on 4th toe.

Tail 113 mm in length, first 72 mm original, last 41 mm regenerated, 9.3 mm in width at base, tapering to a point; dorsal scales of original portion of tail flat, square; original portion segmented, 6–8 transverse scale rows per segment; posterior margin of segments bordered by 3–5 larger tubercles dorsally in anterior one-third of tail, fewer posteriorly; original subcaudal region bearing large, transverse scales; regenerated portion of tail covered with small, smooth to weakly keeled rectangular scales dorsally and laterally and larger scales ventrally; dorsal and lateral caudal furrows extend nearly length of original tail; base of tail bearing hemipenial swellings; four small, postcloacal tubercles on hemipenial swellings; postcloacal scales smooth, flat, large, imbricate.

Coloration in life. Dorsal ground color of head, body, limbs, and tail brown; wide, dark brown, nuchal band extends from posterior margin of one eye to posterior margin of other eye; nuchal band edged with thin, white, broken lines that are most prominent where they encompass tubercles, giving it a somewhat spotted appearance; four similarly colored dorsal bands with similar white edging occur between limb insertions, 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 1.00; one additional dark brown band posterior to hind limbs; no band on posterior margin of thigh; four diffuse, indistinct dark bands and five indistinct lighter bands extend onto original portion of tail; no bands on regenerated portion tail; no caudal or body bands have lighten centers or encompass yellowish to white tubercles; ventral surfaces of head, limbs, and tail smudged with brown; abdomen immaculate, beige except for slightly darker, lateral regions.

Variation. The paratypes closely match the holotype in all aspects of dorsal banding and coloration ( Fig. 10 ). In LSUHC 9124 the dorsal bands are not as distinctly edged with thin white lines and in LSUHC 9123 the third dorsal band bifurcates on the left side at the vertebral region. ZRC 2.6967 has a few light colored tubercles within the lateral portions of the second, third, and fourth bands. ZRC 2.6968 is a gravid female with a slight more opaque overall pattern and there are some yellowish tubercles on the flanks and limbs. A significant degree of ontogenetic change in color pattern occurs with hatchlings and juveniles having a greatly contrasting banding pattern where the body bands are very dark and are edged with thin white lines and the white colored portion of the tail is lost in adulthood. Meristic differences in the type series and additional specimens examined are presented in Table 9.

Additional specimens examined. — Four additional specimens from the type locality were examined ( Table 9) and of these three ( LSUHC 9125, 9435–36) had bifurcations in the third band ( Fig. 10 ). The posterior 50% of the tails in the juveniles LSUHC 9125 and 9437 (SVL 43 mm, 51 mm, and 51 mm, respectively) have a whitish wash ( Fig. 10 ) and appear opaque in life.

Distribution. Cyrtodactylus langkawiensis sp. nov. is known only from Wat Wanaram, Pulau Langkawi, Peninsular Malaysia ( Fig. 3 ).

Natural history. Like its sister species Cyrtodactylus astrum sp. nov., C. langkawiensis sp. nov. is a saxicolous species inhabiting lowland forests in close association with karst formations on which it maybe a substrate specialist ( Fig. 9 ). Lizards were found at night on the sides of karst towers and in cracks from 1–3 meters above the forest floor. None were seen moving about on the ground. Gravid females carrying two eggs were found during October and juveniles were found during March, September, and October indicating C. langkawiensis sp. nov. may breed year round. No C. macrotuberculatus were observed at or near the type locality of C. langkawiensis sp. nov. and surveys throughout much of the forested areas of Pulau Langkawi lacking karst have resulted in the presence of only C. macrotuberculatus . The range of C. langkawiensis sp. nov. is almost certainly not restricted to the type locality and it is expected that this species will be discovered in additional regions with karst formations. Grismer (2011) noted that juveniles bearing white tails wave them over their head when threatened.

Etymology. The specific epithet, langkawiensis , refers to Langkawi Island to where this species was first discovered and may be endemic.

Comparisons. Cyrtodactylus langkawiensis sp. nov. is separated from C. macrotuberculatus in lacking large tubercles on the sp. nov. dorsal surface of the head, body and limbs, on the underside of the forearms, in the gular region, in the ventrolateral body fold; and having 38–43 as opposed to 17–28 ventral scales. From C. pulchellus , C. macrotuberculatus , and C. trilatofasciatus sp. nov., C. langkawiensis sp. nov. differs in having more than 37 ventral scales as opposed to having 36 or less. It differs further from C. australotitiwangsaensis sp. nov., C. trilatofasciatus sp. nov. in having 19–21 as opposed to 21–25 and 22–27 fourth toe lamellae, respectively. Cyrtodactylus langkawiensis sp. nov. has 30 femoro-precloacal pores, separating it from C. astrum sp. nov., C. australotitiwangsaensis sp. nov., C. bintangtinggi sp. nov., C. bintangrendah sp. nov., C. pulchellus and C. trilatofasciatus sp. nov. which have 31 or more pores. It differs further from C. trilatofasciatus sp. nov. in having four or five body bands with a band/interspace ratio of 0.75–1.00 as opposed to having three bands with a ratio of 2.00–2.75. From C. astrum sp. nov. it differs in lacking a scattered dorsal pattern of white dorsal tubercles. Cyrtodactylus langkawiensis sp. nov. differs from all other species except C. astrum sp. nov. and C. lekaguli sp. nov. in that hatchlings and juveniles have white tail tips as opposed to lacking white tail tips. Hatchlings and juveniles of C. trilatofasciatus sp. nov. have not been observed. Having 11–16 dark caudal bands separates it form C. australotitiwangsaensis sp. nov., C. bintangtinggi sp. nov., C. macrotuberculatus , C. pulchellus , and C. trilatofasciatus sp. nov. which have less than 11 dark caudal bands. The heavy infusion of dark mottling in the white caudal bands of adults differentiates C. langkawiensis sp. nov. from C. australotitiwangsaensis sp. nov., C. bintangtinggi sp. nov., C. bintangrendah sp. nov., C. pulchellus , and C. trilatofasciatus sp. nov. which have immaculate white caudal bands. Cyrtodactylus langkawiensis sp. nov. has a maximum SVL of 99.8 mm which separates for all other species, except C. lekaguli sp. nov., that have maximum SVLs greater than 108.3 mm.

Langkawi, Kedah, 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; /=data unavailable.

ZRC ZRC ZRC LSUHC LSUHC ZRC LSUHC LSUHC LSUHC LSUHC 2.6966 2.6967 2.6968 9123 9124 2.6969 9125 9435 9436 9437 Holotype Paratype Paratype Paratype Paratype Paratype

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 9. Cyrtodactylus langkawiensis from Wat Wanaram, Pulau Langkawi, Kedah, Peninsular Malaysia. Upper left: LSUHC 9123, adult female. Upper right: LSUDPC 4694; gravid female. Lower left: LSUDPC 5196; adult male. Lower right: karst microhabitat of C. langkawiensis at Wat Wanaram, Kedah, Peninsular Malaysia. All photos by L. Grismer.

FIGURE 10. Type series and additional specimens examined of Cyrtodactylus langkawiensis. Upper row: type series from left to right; holotype ZRC 2.6966, paratypes from left to right ZRC 2.6967 – 68, LSUHC 9123 – 24, ZRC 2.6969. Lower row: additional specimens examined from type locality from left to right; LSUHC 9436, 9435 and three juveniles.