Cyrtodactylus bintangtinggi, 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 : 28-32

Cyrtodactylus bintangtinggi sp. nov.

Bintang Mountain Bent-toed Gecko Figs. 11 , 12

Gymnodactylus pulchellus Flower 1899:626 (in part); Boulenger 1903:148, 1912:36 (in part); Smith 1930:13. Cyrtodactylus pulchellus Grismer et al. 2010:149, 2011:416 (in part).

Holotype.—Adult male ( ZRC 2.6970) collected on 24 March 2008 by L. Grismer and R. Johnson from Bukit Larut, Peninsular Malaysia (04° 51.715 N, 100°47.993) at 1151 meters above sea level.

Paratypes.—All paratypes come from the same locality as the holotype. Adult male ( ZRC 2.6971) bears the same collection data as the holotype. The adult females ( ZRC 9008 and 9010) and the adult males ( ZRC 2.6972 and LSUHC 9006–07, 9009) were collected on 16 June 2008 by J. Grismer, P. Wood, Jr., R. Gregory, and L. Grismer.

Diagnosis.—Adult males reaching 111.1 mm SVL, adult females reaching 108.3 mm SVL; 9–13 supralabials, 8–11 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; 31–42 paravertebral tubercles; 21–26 longitudinal rows of dorsal tubercles; 36–40 rows of ventral scales; 21–24 subdigital lamellae on fourth toe; 39–41 femoro-precloacal pores in males; dorsum not bearing a scattered pattern of white tubercles; four (rarely three; one of 14 specimens examined) very dark body bands in adults lacking lightened centers and light colored tubercles; band to interspace ratio 1.00–1.25; 8–10 dark caudal bands on original tail; white caudal bands in adults immaculate; 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 104.2 mm; head large, moderate in length (HL/SVL 0.28) and wide (HW/HL 0.72), somewhat flattened (HD/HL 0.37), distinct from neck, and triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; frontal and prefrontal regions deeply concave; canthus rostralis sharply rounded anteriorly; snout elongate (ES/HL 0.44), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.07); ear opening elliptical, moderate in size (EL/HL 0.70), vertically oriented; eye to ear distance greater than diameter of eye; rostral rectangular, divided dorsally by a small postrostral (=internasal) and inverted V-shaped furrow, 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 single postnasal, ventrally by first supralabial; 9(R,L) square supralabials extending to just beyond upturn of labial margin, tapering abruptly below midpoint of eye; first supralabial largest; 9(R,L) infralabials tapering in size posteriorly; scales of rostrum and lores flat, larger than granular scales on top of head and occiput, those on posterior portion of canthus rostralis larger; scales of occiput intermixed with small tubercles; large, boney frontal ridges bordering orbit confluent with boney, V-shaped, 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 45% 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 0.43) 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 five 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 25 longitudinal rows of tubercles at midbody; 35 paravertebral tubercles; 37 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 0.18); scales on dorsal surfaces of forelimbs flat to granular, intermixed with a few 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 well-developed, sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.28), 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; 38 contiguous, pore-bearing femoro-precloacal scales forming an inverted T bearing a deep, precloacal groove in which eight pore-bearing scales are found (four on each 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; 23 (R,L) subdigital lamellae on 4th toe.

Tail 124 mm in length, 9.6 mm in width at base, tapering to a point; dorsal scales of tail flat, squarish; tail segmented with five transverse scale rows per segment; posterior margin of segments bordered by four larger tubercles dorsally in anterior one-third of tail, fewer posteriorly; subcaudal region bearing large, transverse scales; dorsal and lateral caudal furrows extend nearly entire length of tail; base of tail bearing hemipenial swellings; three small, postcloacal tubercles on hemipenial swellings; postcloacal scales smooth, flat, large, imbricate.

Coloration in life ( Fig. 11 ). Dorsal ground color of head, body, limbs, and tail dark brown; faint, white chevron on rostrum; wide, nearly black, nuchal band extends from posterior margin of one eye to posterior margin of other eye; nuchal band edged with prominant thin, white, lines; four similarly colored dorsal bands between limb insertions with immaculate, white, edging most prominent where it encompasses tubercles resulting in a somewhat spotted appearance; 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; seven dark caudal bands at least twice the width of seven white caudal bands extend onto tail; no caudal or body bands have lightened centers or encompass yellowish to white tubercles; ventral surfaces of head, limbs, and tail smudged with brown; gular scales stippled; abdomen immaculate, beige except for slightly darker, lateral regions, and weakly stippled ventral scales.

Variation. The paratypes approximate the holotype in all aspects of dorsal banding and coloration ( Fig. 12 ). LSUHC 8862, 9008, and 9010 lack a white chevron marking on the rostrum. LSUHC 9009 has three dorsal bands. Portions of the tail of LSUHC 8862 and 9008 are regenerated. The white edging of the dark dorsal bands is not as prominent in LSUHC 9006 and it is even less so in LSUHC 9007. LSUHC 9007 is also missing its tail. The color pattern of hatchlings and juveniles has a greatly contrasting banding pattern where the body bands are nearly solid black, only faintly edged in yellow, and the interspaces and top of the head are yellow ( Fig. 11 ). Meristic differences in the type series and additional specimens examined are presented in Table 10 .

Additional specimens examined. —The posterior 50% of the tails in the juveniles LSUDPC 4780 and 5357 were banded not white ( Fig. 11 ).

Distribution. Cyrtodactylus bintangtinggi sp. nov. is known from the type locality of Bukit Larut, Perak and from Gunung Bubu, Perak 20 km to the southwest in Peninsular Malaysia ( Grismer 2011; Grismer et al. 2010a; Fig. 3 ). It is likely, however, this species ranges throughout the Banjaran Bintang farther north into Thailand, similar to the upland Banjaran Bintang endemic Cnemaspis mcguirei ( Grismer et al. 2008b).

Natural history. Cyrtodactylus bintangtinggi sp. nov. is a nocturnal upland species inhabiting hill dipterocarp and montane forests ( Fig. 11 ) ranging up to 1500 m in elevation ( Grismer 2011). It is always found in the vicinity of granite rocks where lizards take refuge in cracks and occasionally beneath rocks on the ground. Lizards can be found climbing on vegetation but only if the vegetation is in the vicinity of rocks. Grismer (2011) reports finding lizards in houses and often occurring in male female pairs. Gravid females carrying two eggs and hatchlings have been found during March ( Grismer 2011).

ZRC ZRC ZRC LSUHC LSUHC LSUHC LSUHC LSUHC ZRC ZRC ZRC ZRC ZRC ZRC Etymology. The word bintangtinggi is a combination of the Malaysian words bintang meaning star and tinggi meaning tall or high and is in reference to the upland regions of the Bintang Mountain Range to where this species is presumably endemic; bintangtinggi is a noun in apposition and is thus invariable.

Comparisons. Cyrtodactylus bintangtinggi 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 in the ventrolateral body fold. From C. macrotuberculatus and C. pulchellus it differs in having more than 35 ventral scales. Having 38–41 femoro-precloacal pores separates it from C. langkawiensis sp. nov. and C. lekaguli sp. nov. which have less than 37 pores and from C. astrum sp. nov. which has 31–38 pores (overlapping by one scale at 38). It differs from C. trilatofasciatus sp. nov. by having four (only one specimen of 14 having three) body bands with a band/interspace ratio of 1.00–1.25 as opposed to having three bands with a band/interspace ratio of 2.00–2.75. Cyrtodactylus bintangtinggi sp. nov. is further separated from C. astrum sp. nov. in lacking a scattered dorsal pattern of white tubercles. From C. astrum sp. nov., C. langkawiensis sp. nov., and C. lekaguli sp. nov., C. bintangtinggi sp. nov. differs in not having hatchlings and juveniles with white tail tips. Cyrtodactylus bintangtinggi sp. nov. is further differentiated from C. astrum sp. nov., C. langkawiensis sp. nov, and C. lekaguli sp. nov. in having fewer than 11 dark caudal bands and from C. trilatofasciatus sp. nov. it differs in having more than seven dark caudal bands. The white caudal bands of C. bintangtinggi sp. nov. are immaculate which differentiates it from C. astrum sp. nov., C. langkawiensis sp. nov., C. lekaguli sp. nov., and C. macrotuberculatus whose white caudal bands are infused with dark pigment. The SVL of 111.1 mm separates C. bintangtinggi sp. nov. from C. australotitiwangsaensis sp. nov., C. macrotuberculatus , and C. trilatofasciatus sp. nov. whose SVLs are greater than 117 mm and it is separated from C. langkawiensis sp. nov. and C. lekaguli sp. nov. whose SVLs are less than 104 mm.

Remarks. Cyrtodactylus bintangtinggi sp. nov. and C. bintangrendah sp. nov. are sister species forming what is referred to here as the Bintang clade. Like other lineages associated with the Banjaran Bintang, the Bintang clade is most closely related to lineages within the Banjaran Titiwangsa, namely the Titiwangsa clade containing the sister species C. trilatofasciatus sp. nov. and C. australotitiwangsaensis sp. nov. This common biogeographic pattern is seen between the bufonids Ansonia malayana Inger and A. jeetsukumarani Wood, Grismer, Norhayati & Senawi ( Wilkinson et al. 2012) , the agamids Acanthosaura bintangensis Wood, Grismer, Grismer, Norhayati, Chan & Bauer and A. titiwangsaensis Wood, Grismer, Grismer, Norhayati, Chan & Bauer (Wood et al. 2009) , the gekkonids Hemiphyllodactylus harterti Werner and H. titiwangsaensis Zug (Zug 2010) , and populations of the colubrids Asthenodipsas vertebralis (Boulenger) (Laredo et al. in prep), Collorhabdium williamsoni Smedley , Lycodon butleri Boulenger , and Macrocalamus chanardi David & Pauwels. The sister lineages are likely to have initially diverged during periods of lowered sea levels that created warming and drying trends in the intervening lowlands and forced cool, more mesic adapted lowland forests to migrate upslope into current montane areas thus bifurcating lowland lineages that initially retreated with the forests to their respective upland refugia.

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 11. Cyrtodactylus bintangtinggi. Upper left: ZRC 2.6970 (holotype); adult male from Bukit Larut, Perak, Malaysia (photo by L. Grismer). Upper right: LSUDPC 5357; hatchling from Bukit Larut, Perak, Malaysia (photo by L. Grismer). Lower left: LSUDPC 6319; adult female from Gunung Bubu, Perak, Malaysia (photo by D. Belabut). Lower right: granite rock microhabitat of C. bintangtinggi at Bukit Larut, Perak, Malaysia (photo by L. Grismer).

FIGURE 12. Type series of Cyrtodactylus bintangtinggi. Upper row from left to right: holotype ZRC 2.6970, paratypes from left to right ZRC 2.6971 – 72, LSUHC 9006. Lower row: paratypes from left to right LSUHC 9007, 9008, 9009, 9010.

Tables

TABLE 10. Morphological data of type series and additional specimens of Cyrtodactylus bintangtinggi form Bukit Larut, Perak, 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.

	2.6970	2.6971	2.6972	9006	9007	9008	9009	9010	2.1174	2.1171	2.5658	2.5659	2.5666	2.4864
	Holotype	Paratype	Paratype	Paratype	Paratype	Paratype	Paratype	Paratype
Sex	m	m	m	m	m	f	m	f	f	m	m	m	m	m
Supralabials	9	12	11	11	10	10	12	12	9	13	11	10	12	12
Infralabials	9	10	8	10	10	9	9	10	10	9	11	10	11	11
Tubercles on ventral surface of forelimbs	0	0	0	0	0	0	0	0	0	0	0	0	0	0
Tubercles in gular region	0	0	0	0	0	0	0	0	0	0	0	0	0	0
Ventrolateral fold tuberculate	0	0	0	0	0	0	0	0	0	0	0	0	0	0
No. of paravertebral tubercles	35	37	31	40	42	39	32	40	35	32	37	34	33	38
No. longitudinal rows of tubercles	25	24	24	21	23	26	24	25	24	24	22	24	25	24
No. of ventral scales	37	40	36	38	39	38	/	39	39	40	36	38	39	26
Proximal subdigital lamellae square	0	0	0	0	0	0	0	0	/	/	0	0	0	0
No.of subdigital lamellae on 4th toe	23	23	23	23	23	24	24	22	/	/	23	21	24	23
No. of femoro- precloacal pores	38	39	40	41	41	/	41	/	/	/	40	37	39	41
Bands on base of thigh	0	0	0	0	0	0	0	0	0	0	0	0	0	0
No. body bands	4	4	4	4	4	4	3	4	4	4	4	4	4	4
Band/interspace width	1.00	1.00	1.25	1.25	1.00	1.25	1.25	1.25	/	/	/	/	/	/
No. dark bands on original tail	8	/	/	8	/	10	/	9	/	/	/	/	/	/
Hatch./juv. with white tail tips	/	/	/	/	/	/	/	/	/	/	/	/	/	/
SVL	104.2	107.8	103.9	100.2	111.1	102.6	104.7	108.3	/	/	/	110.1	/	/
TL	124	113	104	137	/	135	65	137
TW	9.6	10.3	10.1	9	10.4	8.7	10.4	9
FL	18.8	17.3	17.1	15.8	18.8	17.9	16.4	17.5
TBL	20.5	20.8	20.3	19.7	22.4	19.3	20.6	20.4
AG	42.1	47.3	43.6	42.3	46.8	46.4	42.5	49.9
HL	29.3	29.9	30.1	27.8	31.7	29.8	29.8	30.5
HW	21.2	22	21.5	20.3	22.2	20.4	21.4	20.8
HD	10.8	11.2	11.1	11.5	11.6	11.9	11	11.4
ED	6.9	6.7	7	6.4	7.8	6.8	6.5	7.2
EE	8.8	8.8	9.3	8.1	8.8	8.3	8.6	9.3
ES	12.9	12.7	13.7	12.6	15	13.5	13.6	13.6
EN	9.8	8.6	10.1	9.8	11.2	10	9.7	9.8
IO	6.1	5.6	7	7.2	6.9	6.5	6.4	6.3
EL	2.1	2.2	2.6	2	2.7	1.9	2.3	2.2
IN	3.1	3.5	3.4	3.5	3.6	3.2	2.9	3.2