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 Author Grismer, Lee Author Wood, Perry L. Author Quah, Evan S. H. Author Anuar, Shahrul Author Muin, Abdul Author Sumontha, Montri Author Ahmad, Norhayati Author Bauer, Aaron M. text Zootaxa 2012 3520 1 55 journal article 10.5281/zenodo.211847 9ac583a7-c77c-49a4-93a5-dedcd7789325 1175-5326 211847 Cyrtodactylus astrum sp. nov. Starry Bent-toed Gecko Figs. 5 , 6 Cyrtodactylus pulchellus Chan, Grismer, Sharma, Belabut and Norhayati 2009 :280 ; Chan, Grismer, Shahru, Quah, Grismer, Wood, Jr., Muin, & Norhayati 2011 :255 ; Grismer 2011 :415 (in part). Holotype .—Adult male ( ZRC 2.6962) collected on 24 March 2011 by L. L. Grismer, S. Anuar, E. S. H. Quah, Chan, K. O., and M. A. Muin from Wang Kelian, Perlis, Peninsular Malaysia (06° 41.805 N , 100° 10.751 E ) at 150 meters above sea level. Paratypes .— ZRC 2.6963, LSUHC 8810 and 8816 (adult male, female, and male, respectively) collected on 3 March 2008 by L. L. Grismer and Chan K. O. from Gua Kelam, Perlis, Peninsular Malaysia and an adult female ( ZRC 2.6964) and adult male ( ZRC 2.6965) collected on 11 October 2009 and 21 May 2010 , respectively, by E. S. H. Quah, S. Anuar, and M. A. Muin from Gua Wang Burma , Perlis, Peninsular Malaysia (06° 42.005 N , 100° 11.878 E ) at 180 meters in elevation. FIGURE 5. Cyrtodactylus astrum . Upper left: ZRC 2.6962 (holotype); adult male from Wang Kelian, Perlis, Malaysia (photo by L. Grismer). Upper right: LSUDPC 6317; adult male from La-ngu District, Satun Province, Thailand (photo by M. Sumontha). Lower left: LSUHC 10024; hatchling from Wang Kelian, Perlis, Malaysia (photo by L. Grismer). Lower right: karst habitat of C. astrum in Perlis State Park, Perlis (photo by L. Grismer). Diagnosis .—Adult males reaching 108.3 mm SVL, adult females reaching 104.8 mm SVL; 10–12 supralabials, 9–12 infralabials; tubercles of dorsum small to moderately large with several intervening smaller tubercles; no tubercles on ventral surfaces of forelimbs, gular region, or in ventrolateral body fold; 40–57 paravertebral tubercles; 20–29 longitudinal rows of dorsal tubercles; 31–46 rows of ventral scales; 20–24 subdigital lamellae on fourth toe; 31–38 femoro-precloacal pores in males; dorsum bearing a scattered pattern of white tubercles; four dark body bands in adults with lightened centers and light colored tubercles; band to interspace ratio 1.00–2.00; 13 or 14 dark caudal bands on original tail; white caudal bands in adults heavily infused with dark pigmentation, nearly indiscernible; 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 108.3 mm ; head large, moderate in length (HL/SVL 0.39) and wide (HW/HL 0.69), somewhat flattened (HD/HL 0.38), 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 0.45), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.23); ear opening round, moderate in size (EL/HL 0.10), 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 and large 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 large 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,L) infralabials tapering in size posteriorly; 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 largest; 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 65% 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. FIGURE 6. Type series and additional specimens examined of Cyrtodactylus astrum . Upper: Type series from left to right; holotype ZRC 2.6962, paratypes ZRC 2.6963, LSUHC 8810, 8816, ZRC 2.6964, 2.6965. Lower: additional specimens examined. Body relatively short (AG/SVL 0.39) with well-defined, non-tuberculate, ventrolateral folds; dorsal scales small, granular, interspersed with large, trihedral, regularly arranged, keeled tubercles separated by smaller keeled tubercles 50% their size; 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; tubercles absent from regenerated portion of tail; tubercles on occiput and nape relatively small, those on body largest; approximately 20 longitudinal rows of tubercles at midbody; 47 paravertebral tubercles; 46 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 granular, intermixed with 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 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 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; 32 contiguous, pore-bearing femoro-precloacal scales forming an inverted T bearing a deep, precloacal groove in which six pore-bearing scales are found (three 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 rounded; digits welldeveloped, 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 99 mm in length, first 48 mm original, last 51 mm regenerated, 11.4 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 ventrally; dorsal and lateral caudal furrows extend entire length of original tail; base of tail bearing hemipenial swellings; three small, postcloacal tubercles on hemipenial swellings; postcloacal scales smooth, flat, large, imbricate. Coloration in life ( Fig. 5 ). 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, third band bifurcates on right side of body at vertebral column, fourth band terminates on anterior margin of hind limb insertions; body band/interspace ratio 1.50; one additional dark brown band posterior to hind limbs; no band on posterior margin of thigh; diffuse, indistinct bands extend onto original portion of tail becoming less discernable posteriorly; no bands on regenerated portion tail; all bands have light brown centers bearing yellowish to white tubercles, giving dorsum an overall “starry” appearance; 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 and coloration ( Fig. 6 ) except the third body band in the paratypes is not bifurcated. LSUHC 8810 and 8816 have a lighter overall color pattern and LSUHC 8816 has a unicolor regenerated tail. ZRC 2.6964 lacks a tail and its white, dorsal tuberculation is not as prominent. ZRC 2.6965 has a completely regenerated tail and an overall lighter dorsal pattern. ZRC 2.6963 has a slightly more contrasting dorsal pattern. Females (ZRC 2.6964) lack a precloacal groove and pore-bearing femoroprecloacal scales although enlarged homologous scales are present. Overall coloration lightens considerably at night. 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 do not have lighten centers or encompass light colored tubercles. Furthermore, the bands are edged with thin yellow lines, there are no white tubercles in the interspaces, and the top of the head is yellowish ( Fig. 5 ). Lastly the white colored portion of the tail is lost in adulthood. Additional specimens examined. The posterior 50% of the tail in the juveniles LSUHC 8815 and 8808 (SVL 43 mm and 46 mm , respectively) from Gua Kelam and LSUHC 10024 (SVL 46 mm ) from Wang Kelian are white ( Fig. 6 ). Meristic differences in the type series and additional specimens examined are presented in Table 7 . Distribution. Cyrtodactylus astrum sp. nov. ranges from at least La-ngu District, Satun Province, Thailand south to Kuala Perlis, Perlis, Peninsular Malaysia ( Fig. 3 ). Natural history. Cyrtodactylus astrum sp. nov. is a saxicolous species inhabiting lowland forests of varying types and is closely associated with karst formations ( Fig. 5 ). In Perlis, we have found lizards deep within the cave system of Gua Wang Burma as well as on karst walls at Wang Kelian and Kuala Perlis. At Wang Kelian we have collected specimens from the trunks of trees adjacent to karst towers and have seen hatchlings on the ground nearby karst towers during March. In Perlis State Park, C. astrum sp. nov. and C. macrotuberculatus are narrowly syntopic being that we have found both species within Gua Wang Burma . Generally however, these two species partition their habitat in that C. astrum sp. nov. is principally saxicolous but will opportunistically use tree trunks if nearby and C. macrotuberculatus generally occurs on vegetation but will opportunistically use rock surfaces (including karst) if nearby ( Grismer 2011 ). Lizards observed in the La-ngu District of Satun Province, Thailand were also associated with karst formations. Cyrtodactylus lekaguli sp. nov. occurs at this locality as well (see below). Grismer (2011) noted that juveniles bearing white tails will wave them over their head when threatened. Etymology. The specific epithet astrum is Latin, meaning “a star” and is in reference to the starry pattern of the white dorsal tubercles. TABLE 7. Morphological data of type series and additional specimens of Cyrtodactylus astrum from Perlis 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. B = broken; PR = partially regenerated; R = regenerated; /=data unavailable.

LSUHC	ZRC	LSUHC	ZRC	ZRC	ZRC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC
8816	2.6963	8810	2.6964	2.6962	2.6965	8806	8808	8815	9125	10024
Sex Supralabials	Paratype Kuala Perlis m 12	Paratype Gua Kelam m 12	Paratype Gua Kelam f 12	Paratype Kuala Perlis f 11	Holotype Wang Kelian m 12	Paratype Gua Wang Burma m 10	Gua Kelam / 10	Gua Kelam / 12	Gua Wang Burma / 10	Gua Wang Burma / 12	Wang Kelian / 11
Infralabials Tubercles on ventral surface of forelimbs	10 0	10 0	9 0	12 0	10 0	10 0	9 0	10 0	9 0	9 0	10 0
Tubercles in gular region	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
No. of paravertebral tubercles	57	43	45	50	47	45	45	48	40	44	45
No. longitudinal rows of tubercles No. of ventral scales	26 31	22 41	22 39	23 42	20 46	22 41	28 38	23 41	29 37	23 38	22 41
Proximal subdigital lamellae square No. of subdigital lamellae on 4th toe	0 21	0 20	0 22	0 24	0 20	0 21	0 23	0 24	0 22	0 23	0 22
No. of femoro- precloacal pores	38	32	/	/	32	31	/	/	/	/	/
Bands on base of thigh No. body bands	0 4	0 4	0 4	0 4	0 4	0 4	0 4	0 4	0 4	0 4	0 4
Band/interspace width	1	1	1	2.00	1.50	1	1	1	1	1.50	1
No. dark bands on original tail	/	/	/	/	/	/	/	/	/	13	14
Hatch./juv. with white tail tips	/	/	/	/	/	/	Yes	Yes	/	Yes	Yes
SVL TL	97.1 91.8/R	97.9 109/PR	92.2 60.2/B	104.8 B	108.3 99/PR	103.7 102/R	46.4	45.6	42.5	50.0	46.1
TW	8.7	10.4	8.1	8.8	11.4	10.7
FL TBL AG	17 19.2 37.4	17.4 20.6 38.4	16 18.6 36.6	17.6 20.8 45.6	17.3 21.8 42.5	18.5 22.1 43.7
HL HW HD	28.3 19.7 10.8	28.1 19.4 11	26.5 18.3 10.5	30.1 20.5 11.3	30.7 21.3 11.8	30 20.5 12.1
ED EE ES	6.9 8.6 12.7	7.7 7.7 12.7	6.2 7.38 11.6	6.5 8.4 13.3	7.1 8.7 13.7	6.6 8.2 13.2
EN IO EL	9.7 6.3 2.5	9.4 5.8 2.7	10 5.7 2.8	10.5 6.3 2.4	10.1 7 3	10.1 6.3 2.3
IN	2.8	2.7	2.8	3	3	2.9

Comparisons. Cyrtodactylus astrum sp. nov. is separated from all other members of the Cyrtodactylus pulchellus complex by its scattered pattern of white, dorsal tubercles. From C. macrotuberculatus it differs 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; and having 31–46 ventral scales as opposed to 17–28 ( Table 6 ). It differs from C. trilatofasciatus sp. nov. in having more than 39 paravertebral tubercles as opposed to having 38 tubercles or less. It differs further from C. langkawiensis sp. nov. and C. lekaguli sp. nov. in having dark body bands in adults with lightened centers as opposed to these bands being unicolor. From C. trilatofasciatus sp. nov. it differs in having four as opposed to three dark dorsal bands and in having a body band/interspace ratio of 1.00–2.00 vs. 2.00–2.75. It differs from all other species except C. langkawiensis sp. nov. and C. lekaguli sp. nov. in that hatchlings and juveniles have white tail tips as opposed to lacking white tail tips. Cyrtodactylus astrum sp. nov. has 13 or 14 dark caudal bands on the original tail which differentiates it from C. australotitiwangsaensis sp. nov. , C. bintangtinggi sp. nov. , C. pulchellus , C. trilatofasciatus sp. nov. and C. macrotuberculatus which have less than 11 caudal bands. Heavy dark mottling in the white caudal bands differentiate it from all other species except C. langkawiensis sp. nov. , C. lekaguli sp. nov. , and C. macrotuberculatus . The maximum SVL of 108.3 mm of C. astrum sp. nov. separates it from C. australotitiwangsaensis sp. nov. , C. bintangrendah sp. nov. , C. macrotuberculatus , C. pulchellus and C. trilatofasciatus sp. nov. whose maximum SVL is greater than 114.0 mm and from C. langkawiensis sp. nov. and C. lekaguli sp. nov. whose maximum SVL is less than 103.6 mm . Remarks. The sister species relationship between Cyrtodactylus astrum sp. nov. and C. langkawiensis sp. nov. ( Fig. 1 ) is in biogeographical accordance with that of other species found south of the Isthmus of Kra that range south into the Banjaran Nakawan along the northwestern portion of the Malaysian border and whose sister lineage occurs on the offshore extension of this range, Pulau (=island) Langkawi. Such taxa include Cnemaspis roticanai ( Grismer 2011 ) and Cryptelytrops venustus (Grismer et al . in prep.).